Coral reef resilience annotated bibliography Abelson, A, Olinky, R and Gaines, S. (2005) Coral recruitment to the reefs of Eilat, Red Sea: Temporal and spatial variation, and possible effects of anthropogenic disturbances. Marine Pollution Bulletin 50, 576-582. The accelerating deterioration of the coral reefs of Eilat has raised debate over the exact causes and how they affect the reefs. The hypothesis of the present study was that a low recruitment rate of reef-building coral species may play an important role in the decline of the Eilat reefs. Our goal was to assess spatial and temporal recruitment patterns in Eilat, focusing on examining the possible impact of human activities. The results of coral recruitment to 10 series of ceramic tiles on metal racks, revealed very low overall recruitment relative to other geographical regions. In addition, we found that recruitment rates and recruit survival were lowest at sites closest to the major eutrophication sources in Eilat. The low recruitment rates may be chronically too low to compensate for the elevated coral mortality rates of recent years. The significant differences between the present study and a similar study carried out during the same period using a different method, emphasize the crucial need for a standardized method for recruitment assessment in coral reefs worldwide. Abramovitch-Gottlib, L, Katoshevski, D and Vago, R. (2002) A computerized tank system for studying the effect of temperature on calcification of reef organisms. Journal of Biochemical and Biophysical Methods 50, 245-252. Mediated by algal symbionts, calcification in reef building corals is one of the important processes, which enable coral’s growth. In the present study, we used a buoyant weighing technique to study calcification of two coralline species, Stylophora pistillata and the hydrocoral Millepora dichotoma. The colonies were grown in a tank system, in which light, nutrition and water motion were kept constant and temperature was elevated by means of a computerized controlled apparatus. An almost constant rate of calcification was observed in the two species at 22–28°C. Elevation of the temperature above this range to 29–31°C caused a slow down in calcification in both species. A grater number of S. pistillata colonies became bleached at temperatures of ≥29°C, whereas M. dichotoma colonies suffered from bleaching only after three days at 31°C. For both species, control groups, remained viable during the experimental period. The differences in responses to changes in temperature of the two species may be as a consequence of different adaptive mechanisms or to different susceptibilities of the corals to elevated temperatures. We have shown that elevating temperatures above annual maximal ranges have a significant effect on coral calcification. We also demonstrated that sessile calcified marine organisms having ecological and biomedical significance could be cultured and manipulated under laboratory conditions.

Adger, WN, Hughes, TP, Folke, C, Carpenter, SR and Rockström, J. (2005) Social-ecological resilience to coastal disasters. Science 309, 1036-1039. Social and ecological vulnerability to disasters and outcomes of any particular extreme event are influenced by buildup or erosion of resilience both before and after disasters occur. Resilient social-ecological systems incorporate diverse mechanisms for living with, and learning from, change and unexpected shocks. Disaster management requires multilevel governance systems that can enhance the capacity to cope with uncertainty and surprise by mobilizing diverse sources of resilience. Adjeroud, M, Penin, L and Carroll, A. (2007) Spatio-temporal heterogeneity in coral recruitment around Moorea, French Polynesia: Implications for population maintenance. Journal of Experimental Marine Biology and Ecology 341, 204–218. The spatio-temporal variability of scleractinian coral recruitment was investigated from December 2000 to December 2003 around Moorea Island (French Polynesia). Nine stations, each with 20 terracotta tiles, were placed on the outer reef slope, at 3 sites (Vaipahu, Tiahura, Haapiti), and at 3 depths (6, 12 and 18 m). The relative contribution of the different families of recruits (Pocilloporidae: 60.4%, Poritidae: 18.8%, Acroporidae: 11.2%) and the very low recruitment rates (maximum: 35 recruits per tile, higher mean densities of 10.8 recruits per tile, average 40.77 recruits m−2 year−1) recorded at Moorea are similar to recruitment patterns recorded on sub-tropical reefs. Over the duration of the study period, we detected a marked seasonal variability in recruitment rates, with the peak recruitment for all families recorded in December–March periods, corresponding to periods of warmest SSTs. However, recruits of Acroporidae were also relatively abundant in the September–December period in some years, which coincides with the known spawning periods of some Acropora species. Total recruitment rate decreased after the first year of the survey, and was probably the result of the bleaching event that occurred in early 2002, which may have reduced fecundity of some coral populations. A lower proportion of recruits were found on the upper surface of the tiles (14.5%), compared to the lower surfaces (57.1%) and sides (28.4%), which is likely the result of intense grazing by herbivorous fish and urchins.We detected a patchy distribution at the station scale, and a significant variation in recruitment patterns among depths and sites. Pocilloporidae recruited more abundantly at 6 and 18 m, whereas Poritidae were generally more abundant at 12mdepth. In contrast, Acroporidae showed no clear depth pattern during the study period. Recruitment was lowest at the most exposed site (Haapiti), especially for Acroporidae, and probably reflects lower settlement rates and/or higher early post-settlement mortality caused by frequent high swells and their associated strong currents. These distinctive characteristics in recruitment patterns of the 3 dominant families of recruits underline the important role of life history strategies in understanding the spatial patterns, dynamics and maintenance processes of coral populations. The relatively low coral recruitment rates recorded from this study indicate that recovery from severe or frequent perturbations will be slow.

Ahmed, M, Chong, CK and Cesar, H (eds.) (2004) Economic valuation and policy priorities for sustainable management of coral reefs. WorldFish Centre: Penang (Malaysia), 222 pp. Having been brought more closely to the attention of researchers worldwide by the International Year of the Reef (1997), the issue of conservation of coral reefs has intensified. Stresses on coral reefs created by increased development and population adjacent to the coast call for relevant authorities to take immediate action to prevent additional irrecoverable damage occurring worldwide. As of 2001 it was estimated that 11 per cent of all coral reefs had been totally destroyed or damaged beyond recovery, another 16 per cent had been severely damaged in 1998 by coral bleaching related to climate change.1 Since the workshop that formed the basis of these proceedings, the Climate Prediction Center of the United States has reported that warmer than normal sea surface and subsurface temperatures were observed throughout most of the equatorial Pacific during April 2002. Sea surface temperature anomalies were up to 2°C warmer than average in the region between the Galapagos Islands and the South American coast, and more than 1°C warmer than average immediately to the west of 180°W. The Climate Prediction Center also forecast a slow evolution towards El Niño conditions throughout the remainder of 2002.2 The Australian Bureau of Meteorology, in a comparison study, reported that seven out of 12 reputable ocean or coupled ocean/atmosphere forecast models predicted “warm” temperatures from April to September 2002.3 Such climate change would result in more coral reefs being destroyed. Furthermore, Talbot and Wilkinson (2001) concluded that, largely as a result of locally based rather than natural global stresses, a further 30 per cent of the world’s reefs will be seriously damaged in the next 20 to 40 years. Papers from the WorldFish workshop published in this report suggest that reef damage caused by human impacts needs to be addressed at local, national, regional and global levels. Coral reefs can be sustainably managed if reef uses are optimized and good policies are in place. This volume is the outcome of the “International Consultative Workshop for Economic Valuation and Policy Priorities for Sustainable Management of Coral Reefs” held at the WorldFish Center’s Headquarters, Penang, Malaysia, 8-10 December 2001. The overall goal of the workshop was to identify future economic and policy research directions relevant to the sustainable management of coral reefs. The directions were to be identified through review and discussion of the effectiveness of policy instruments; analysis of past research findings; and analysis of the interdependency of community livelihood, coral reefs and their resources. For more effective policy instruments to be introduced by any government, we believe that economic valuation and cost benefit analysis are important processes. They will provide information on the various values of coral reefs, which could allow decision-makers to devise policies that optimize the services and functions provided by the reef ecosystems and their capacity to support the livelihood of coastal communities. The workshop was the final activity of the Valuation and Policy Analysis for Sustainable Management of Coral Reefs project sponsored by the Center’s donors and the Swedish International Development Cooperation Agency (Sida), with additional support from the International Coral Reef Action Network (ICRAN), and support for selected participants

from Southeast Asia by the Economy and Environment Program for Southeast Asia (EEPSEA) and the Australian Center for International Agricultural Research (ACIAR). A total of 48 participants from 15 countries located in Southeast and East Asia, the Caribbean, East Africa and the South Pacific Regional Seas attended the workshop. Seven keynote papers and 19 research papers were presented at the workshop. These proceedings are organized into four sections and three Appendices. The Introduction – the first section – gives a brief account of current issues and problems in coral reef management. The second section focuses on the economic valuation and socioeconomics of coral reefs and consists of two parts. Part A is made up of four keynote papers which provide an overview of the theory and practice of economic valuation and the socioeconomics of coral reefs; the role of economic valuation in coral reef management over the next decade; the use of modeling as a tool to estimate the economic values of coral reefs; and the need for and potential role of economic valuation in relation to coral reef use and management in the Pacific region. Part B of this section consists of four case studies relating to Malaysia, Tanzania (Zanzibar), Thailand, and Vietnam. Section 3 relates to policy instruments and management techniques for coral reefs and marine resources – this section has two parts. Part A consists of three papers addressing the effectiveness of various policy instruments for coral reefs and fisheries management. Part B consists of case studies of policy implementation, climate change adaptation strategies, and coral reef management in the small and low-lying states of the Caribbean, China, Indonesia and Jamaica. The final section of these proceedings consists of outcomes from the working group discussions. It sets out participants’ recommendations on the preferable directions for future research. Recommendations are grouped into research relating to the economic valuation of coral reefs, coral reef policy analysis, and community participation in coral reef management. The Appendices provide a list of workshop participants and their contact details, the workshop program, and an explaination of abbreviations and accronyms used. The editors would like to thank Sida and ICRAN for their support. We would also like to thank EEPSEA and ACIAR for sponsoring participants from Vietnam and Indonesia, respectively. Special thanks are due to all the keynote presenters who provided a strong background for the working group discussions. Thanks also to the case study presenters who generously shared their experience and research findings in the workshop. Last, but not least, sincere thanks to Dr. Peter Gardiner for his generous time and valuable comments on the first draft of these proceeding

Ammar, MSA, Amin, EM, Gundacker, D and Mueller, WG. (2000) One rational strategy for restoration of coral reefs: Application of molecular biological tools to select sites for rehabilitation by asexual recruits. Marine Pollution Bulletin 40, 618-627. Experiments for reef rehabilitation were performed at two selected sites near Hurghada (Red Sea, Egypt) the reef close to the Marine Biological Station (with a high sedimentation rate from landfilling) and El-Fanadir Reef (a clear water site). Since only little is known about the influence of the physical environmental conditions, novel molecular biological approaches have been introduced to assess the metabolic status of corals. In order to avoid possible interference with symbionts the molecular studies have been performed with the octocoral Dendronephthya klunzingeri; this species does not contain zooxanthellae. The metabolic enzymes fructose-1,6-bisphosphatase and the succinat-dehydrogenase served as markers for the assessment of the health status of the corals. The cDNAs for both enzymes were isolated and their levels of expression were found to be correlated with the degree of environmental stress. High expression was found at the El-Fanadir Reef, while only low levels were measured at the Marine Biological Station, which is characterized by high sedimentation rates. From this it is concluded that the health state of D. klunzingeri from El-Fanadir is superior to the one from the Marine Biological Station. Six reef-building corals have therefore been selected from El-Fanadir for the transplantation studies. We applied fixation of coral nubbins in plastic meshes with narrow openings. The asexual recruits remained either unfixed or had been glued to the mesh with epoxy resin. A total of 236 coral fragments were transplanted at the Marine Biological Station Reef and 108 fragments at El-Fanadir Reef. The referred technique was successful and the survival rates were higher for samples fixed with epoxy resin than for those without epoxy resin. The survival and growth rates of the coral transplants were found to be higher at the windward side of El-Fanadir Reef than on the leeward side of the same reef. Furthermore, the mortality rates at the leeward side of El-Fanadir Reef were still lower compared to the Marine Biological Station. The coral species Pocillopora damicornis grew well in the clearer water and hard rocky substrate but it did not grow at all in turbid water and sandy substrate; however, the species Acropora humilis grew well in both environments but with higher rates in the clearer water. After one year of transplantation, the massive coral Favia stelligera recorded the highest survival rate of all coral species at the Marine Biological Station Reef; but among branching corals, A. humilis had the highest value and P. damicornis the lowest. Contrary to this result, P. damicornis recorded the highest value at El-Fanadir Reef. It is concluded that asexual recruits of corals, taken from a site (D. klunzingeri from El-Fanadir) physiologically favourable for them, are suitable for a coral restoration strategy.

Anderson, S, Zepp, R, Machula, J, Santavy, D, Hansen, L and Mueller, E. (2001) Indicators of UV exposure in corals and their relevance to global climate change and coral bleaching. Human and Ecological Risk Assessment 7, 1271-1282.

A compelling aspect of the deterioration of coral reefs is the phenomenon of coral bleaching. Through interactions with other factors such as sedimentation, pollution, and bacterial infection, bleaching can impact large areas of a reef with limited recovery, and it might be induced by a variety of stressors including temperature and salinity extremes, and ultraviolet light. Under conditions of ocean warming, often associated with calm and stratified waters, photobleaching of UV-absorbing chromophoric dissolved organic matter (CDOM) is increased, and penetration of both UV-B and UV-A is greatly enhanced. Indices of UV-specific effects in coral tissue are needed to test whether UV increases, associated with global climate change, are harmful to corals. To address this challenge, we have evaluated UV-specific effects in corals and have characterized factors that alter penetration of UV radiation over coral reefs. An immunoblotting assay was developed to examine UV-specific lesions (thymine dimers) in coral and zooxanthellae DNA. We observed dose-dependent increases of thymine dimers in coral (Porites porites var porites) exposed to artificial solar irradiance in a solar simulator, although effects were not strictly proportional. UV measurements were made in July 1999 at Eastern Sambo reef and nearby sites, including profiling along transects from reef to shore. Results of these analyses indicate that the coral at Eastern Sambo reef (at 3-4 meters) were receiving UV-B radiation that was equivalent to 25 to 30% of surface UV irradiance. However, the water just inside the reef in Hawk Channel (located closer to land) was considerably more opaque to UV. This water photobleached with loss of UV absorbance and fluorescence when it was exposed to simulated solar radiation. These results indicate that photobleaching of the DOM and transport of near-shore water out over the reefs might play a key role in controlling UV penetration to the reef surface Anthony, KRN and Kerswell, AP. (2007) Coral mortality following extreme low tides and high solar radiation. Marine Biology 151, 1623-1631. Extreme tidal events are one of the most predictable natural disturbances in marine benthic habitats and are important determinants of zonation patterns in intertidal benthic communities. On coral reefs, spring low tides are recurrent disturbances, but are rarely reported to cause mass mortality. However, in years when extremely low tides coincide with high noon irradiances, they have the potential to cause widespread damage. Here, we report on such an event on a fringing coral reef in the central Great Barrier Reef (Australia) in September 2005. Visual surveys of colony mortality and bleaching status of more than 13,000 corals at 14 reef sites indicated that most coral taxa at wave-protected sites were severely affected by the event. Between 40 and 75% of colonies in the major coral taxa (Acropora, Porites, Faviidae, Mussidae and Pocilloporidae) were either bleached or suVered partial mortality. In contrast, corals at waveexposed sites were largely unaffected (<1% of the corals were bleached), as periodic washing by waves prevented desiccation. Surveys along a 1–9 m depth gradient indicated that high coral mortality was confined to the tidal zone. However, 20–30% of faviid colonies were bleached throughout the depth range, suggesting that the increase in benthic irradiances during extreme low tides caused light stress in deeper water. Analyses of an 8-year dataset of tidal records for

the area indicated that the combination of extended periods of aerial exposure and high irradiances occurs during May–September in most years, but that the event in September 2005 was the most severe. We argue that extreme low-tide, high-irradiance events are important structuring forces of intertidal coral reef communities, and can be as damaging as thermal stress events. Importantly, they occur at a time of year when risks from thermal stress, cyclones and monsoon-associated river run-off are minimal. Apprill, AM, Bidigare, RR and Gates, RD. (2007) Visibly healthy corals exhibit variable pigment concentrations and symbiont phenotypes. Coral Reefs 26, 387-397. Understanding the natural variability of photosynthetic pigment ranges and distributions in healthy corals is central to evaluating how useful these measurements are for assessing the health and bleaching status of endosymbiotic reef-building corals. This study examined the photosynthetic pigment variability in visibly healthy Porites lobata and Porites lutea corals from Kaneohe Bay, Hawaii and explored whether pigment variability was related to the genetic identity or phenotypic characteristics of the symbionts. Concentrations of the photosynthetic pigments chlorophyll a, peridinin, chlorophyll c2, diadinoxanthin, diatoxanthin, β,β -carotene and dinoxanthin were quantiWed using highperformance liquid chromatography (HPLC). Pigment concentrations were found to range 1.5–10 fold in colonies of each species at similar depths (0–2, 2–4, 10–15 and 19–21 m). Despite the high pigment variability, pigment ratios for each species were relatively conserved over the 0–21 m depth gradient. The genetic identity of the symbiont communities was examined for each colony using 18S nuclear ribosomal DNA (nrDNA) restriction fragment length polymorphisms. All colonies contained symbionts belonging to clade C. The density and phenotypic characteristics of the symbionts were explored using Flow cytometry, and Fluorescence and side scatter (cell size) properties revealed phenotypically distinct symbiont subpopulations in every colony. The symbiont subpopulations displayed pigment trends that may be driven by acclimatization to irradiance microenvironments within the host. These results highlight the biological complexity of healthy coral–symbiont associations and the need for future research on pigments and symbiont subpopulation dynamics. Aronson, RB, Precht, WF, Toscano, MA and Koltes, KH. (2002) The 1998 bleaching event and its aftermath on a coral reef in Belize. Marine Biology 141, 435-447. Widespread thermal anomalies in 1997–1998, due primarily to regional effects of the El Niño–Southern Oscillation and possibly augmented by global warming, caused severe coral bleaching worldwide. Corals in all habitats along the Belizean barrier reef bleached as a result of elevated sea temperatures in the summer and fall of 1998, and in fore-reef habitats of the outer barrier reef and offshore platforms they showed signs of recovery in 1999. In contrast, coral populations on reefs in the central shelf lagoon died off catastrophically. Based on an analysis of reef cores, this was the first bleaching-induced mass coral mortality in the central lagoon in at least the last 3,000 years. Satellite data for the Channel Cay reef complex, the most intensively studied of the lagoonal reefs, revealed a prolonged period of elevated sea-surface temperatures (SSTs) in the late summer and early fall of 1998. From 18

September to 1 October 1998, anomalies around this reef averaged +2.2°C, peaking at 4.0°C above the local HotSpot threshold. In situ temperature records from a nearby site corroborated the observation that the late summer and early fall of 1998 were extraordinarily warm compared to other years. The lettuce coral, Agaricia tenuifolia, which was the dominant occupant of space on reef slopes in the central lagoon, was nearly eradicated at Channel Cay between October 1998 and January 1999. Although the loss of Ag. tenuifolia opened extensive areas of carbonate substrate for colonization, coral cover remained extremely low and coral recruitment was depressed through March 2001. High densities of the sea urchin Echinometra viridis kept the cover of fleshy and filamentous macroalgae to low levels, but the cover of an encrusting sponge, Chondrilla cf. nucula, increased. Further increases in sponge cover will impede the recovery of Ag. tenuifolia and other coral species by decreasing the availability of substrate for recruitment and growth. If coral populations are depressed on a long-term basis, the vertical accretion of skeletal carbonates at Channel Cay will slow or cease over the coming decades, a time during which global-warming scenarios predict accelerated sea-level rise. Aronson, RB, Macintyre, IG, Wapnick, CM and O’Neill, MW. (2004) Phase shifts, alternative states, and the unprecedented convergence of two reef systems. Ecology 85, 1876-1891. Initial conditions can generate differences in the biotic composition of spatially disjunct communities, but intense, large-scale perturbations have the potential to reduce or eliminate those historical differences. The latter possibility is of particular concern with respect to coral reefs, which have undergone dramatic changes in the last 25–30 years. This paper reports a case in which two reef systems with different biotic histories were recently perturbed to a single, novel state. We compared millennial-scale records of species dominance from reefs in Bahía Almirante, a coastal lagoon in northwestern Panama, to previously published records from reefs in the shelf lagoon of Belize. Reef cores extracted from Bahía Almirante at 5–10 m water depth revealed that the Panamanian reefs were persistently dissimilar from the Belizean reefs for at least 2000–3000 years prior to the last several decades. The Panamanian reefs were dominated continuously by branching finger corals, Porites spp. (primarily P. furcata). Shifts from the Porites-dominated state to dominance by other coral species were rare, were restricted to small areas, and lasted for decades to centuries. The Belizean reefs were dominated continuously by the staghorn coral Acropora cervicornis in the same depth range during the same period. Excursions from the Acropora-dominated state were again rare and spatially localized. Populations of Ac. cervicornis in the Belizean lagoon were nearly extirpated by an outbreak of whiteband disease in the late 1980s, and changes in water quality were apparently detrimental to branching Porites in Bahía Almirante in recent decades. These large-scale perturbations caused the two reef systems to converge on a third, historically unprecedented state: dominance by the lettuce coral Agaricia tenuifolia. Ag. Tenuifolia possesses life-history attributes and environmental tolerances that enabled it to become dominant in both disturbed ecosystems. Although the two phase shifts to Ag. tenuifolia differed in both their general mechanisms and specific causes, they had the effect of eliminating the salient difference in benthic composition between the Panamanian and Belizean reefs. The changes in species composition thus obliterated the influence of several thousand years of reef history.

Aronson, RB and Precht, WF. (2006) Conservation, precaution, and Caribbean reefs. Coral Reefs 25, 441–450. Some authors argue that overfishing is an important reason that reef corals have declined in recent decades. Their reasoning is that overfishing removes herbivores, releasing macroalgae to overgrow and kill the corals. The evidence suggests, however, that global climate change and emergent marine diseases make a far greater contribution to coral mortality, and that macroalgae generally grow on the exposed skeletal surfaces of corals that are already dead. Macroalgal dominance, therefore, is an effect rather than a cause of coral mortality. Marine protected areas (MPAs), which are usually established to protect stocks of reef fish, foster populations of herbivorous fish under at least some circumstances. Increased herbivory can reduce algal cover, potentially accelerating the recovery of coral populations inside MPAs; however, establishing MPAs will have only a limited impact on coral recovery unless policymakers confront the accelerating negative effects of the global-scale sources of coral mortality. Arthur, R. (2000) Coral bleaching and mortality in three Indian Ocean reef regions during an El Niño Southern Oscillation event. Current Science 79, 1723-1729. The 1997–1998 El Niño Southern Oscillation (ENSO) event, which elevated Sea Surface Temperatures (SSTs) of tropical oceans by more than 3°C, was one of the most extreme ENSO events in recent history. Such increases in SSTs above the seasonal average can trigger widespread bleaching in coral reefs. This study examined bleaching in three Indian coral reef regions in relation to SSTs using quantitative rapid assessment methods between April and July, 1998. The Gulf of Kutch reefs showed an average of 11% bleached coral with no apparent bleaching-related mortality. In contrast, bleached coral comprised 82% of the coral cover in lagoon reefs of Lakshadweep and 89% of the coral cover in the Gulf of Mannar reefs. Bleaching-related mortality was high – 26% in Lakshadweep and 23% in Mannar. The coral mass mortality may have profound ecological and socio-economic implications and highlights the need for sustained monitoring for coral reef conservation in India. Arthur, R, Done, TJ, Marsh, H and Harriott, V. (2006) Local processes strongly influence post-bleaching benthic recovery in the Lakshadweep Islands. Coral Reefs 25, 427–440. The atoll reefs of the Lakshadweep, in the Indian Ocean suffered a catastrophic mortality of hard coral in the wake of the El Niño event of 1998. This study tracked changes to coral and other benthic elements in three atolls in the Lakshadweep from 2000 to 2003. The recovery of coral was highly site-specific, and appeared to be driven by differences in post-settlement survival of coral recruits, that were in turn, influenced by the local hydrodynamics of the atolls. Post bleaching recovery was highest on west-facing reefs, while recovery on east-facing reefs was very limited. However, no ‘phase-shift’ to macroalgal dominated reefs was evident. High herbivore pressures were perhaps the most important control of macroalgae. Five years after the mass mortality, the genera that showed the maximum gains represented a mix of different susceptibilities to bleaching, while some genera that were not particularly

susceptible to bleaching showed significant declines. These results suggest that decline or recovery of coral is likely dependent on individual life history strategies, post-recruitment survival, and contingency. Aw, M. (2000) New wave coral reef rejuvenation – The Ihuru Island Barnacle Reef project. Asian Geographic, 50-57. The hole in the ozone layer, El Niño and global warming are household names today – phenomena with which we are all familiar and which are known to be playing havoc with the natural environment. One effect believed to be caused by such phenomena is the gradual rising of sea temperatures. The devastating effect that rising sea temperatures can have on coral reefs was witnessed in the Maldives in 1998 when a sudden spike in sea temperature inflicted significant damage to what were some of the most magnificent coral reefs of the world. While scientific opinion remains divided as to exactly what caused the temperature spike, its effects were immediately visible even to non-scientists. The good news is that today the reefs in the Maldives are well on their way to recovery and there is strong hope that new technologies currently being tested in the Maldives will play an important role in assisting the rejuvenation of the coral reef systems. Ayre, DJ and Hughes, TP. (2004) Climate change, genotypic diversity and gene flow in reef-building corals. Ecology Letters 7, 273-278. In the ocean, large-scale dispersal and replenishment by larvae is a key process underlying biological changes associated with global warming. On tropical reefs, coral bleaching, degradation of habitat and declining adult stocks are also likely to change contemporary patterns of dispersal and gene flow and may lead to range contractions or expansions. On the Great Barrier Reef, where adjacent reefs form a highly interconnected system, we use allozyme surveys of c. 3000 coral colonies to show that populations are genetically diverse, and rates of gene flow for a suite of five species range from modest to high among reefs up to 1200 km apart. In contrast, 700 km further south on Lord Howe Island, genetic diversity is markedly lower and populations are genetically isolated. The virtual absence of long-distance dispersal of corals to geographically isolated, oceanic reefs renders them extremely vulnerable to global warming, even where local threats are minimal. Bachok, Z, Mfilinge, P and Tsuchiya, M. (2006) Characterization of fatty acid composition in healthy and bleached corals from Okinawa, Japan. Coral Reefs 25, 545-554. Under bleaching conditions, corals lose their symbiotic zooxanthellae, and thus, the ability to synthesize fatty acids (FAs) from photosynthetically derived carbon. This study investigated the lipid content and FA composition in healthy and bleached corals from the Odo reef flat in Okinawa, southern Japan, following a bleaching event. It was hypothesized that the FA composition and abundance would change as algae are lost or die, and possibly microbial abundance would increase in corals as a consequence of bleaching. The lipid content and FA composition of three healthy coral species (Pavona frondifera, Acropora pulchra, and Goniastrea aspera) and of partially bleached and completely bleached colonies of P. frondifera were examined. The FA composition did not differ among healthy corals, but differed significantly

among healthy, partially bleached, and completely bleached specimens of P. frondifera. Completely bleached corals contained significantly lower lipid and total FA content, as well as lower relative amounts of polyunsaturated FAs and higher relative amounts of saturated FAs, than healthy and partially bleached corals. Furthermore, there was a significantly higher relative concentration of monounsaturated Fas and odd-numbered branched FAs in completely bleached corals, indicating an increase in bacterial colonization in the bleached corals. Baghdasarian , G and Muscatine, L. (2000) Preferential Expulsion of Dividing Algal Cells as a Mechanism for Regulating Algal-Cnidarian Symbiosis. Biological Bulletin 199(3), 278-286. A wide range of both intrinsic and environmental factors can influence the population dynamics of algae in symbiosis with marine cnidarians. The present study shows that loss of algae by expulsion from cnidarian hosts is one of the primary regulators of symbiont population density. Because there is a significant linear correlation between the rate of algal expulsion and the rate of algal division, factors that increase division rates (e.g., elevated temperature) also increase expulsion rates. Additionally, 3H-thymidine is taken up to a greater extent by algae destined to be expelled than by algae retained in the host cnidarians. Taken together, data for rates of expulsion, rates of division at different temperatures, and uptake of 3H-thymidine suggest that dividing algal cells are preferentially expelled from their hosts. The preferential expulsion of dividing cells may be a mechanism for regulation of algal population density, where the rate of expulsion of algae may be an inverse function of the ability of host cells to accommodate new algal daughter cells. This kind of regulation is present in some cnidarian species (e.g., Aiptasia pulchella, Pocillopora damicornis), but not in all (e.g., Montipora verrucosa, Porites compressa, and Fungia scutaria). Baird, AH and Marshall, PA. (2002) Mortality, growth and reproduction in scleractinian corals following bleaching on the Great Barrier Reef. Marine Ecology Progress Series 237, 133-141. Despite extensive research into the coral bleaching phenomena there are very few data which examine the population biology of affected species. These data are required in order to predict the capacity of corals to respond to environmental change. We monitored individual colonies of 4 common coral species for 8 mo following historically high sea-surface temperatures on the Great Barrier Reef in 1998 to compare their response to, and recovery from, thermal stress and to examine the effect of bleaching on growth and reproduction in 2 Acropora species. Platygyra daedalea and P. lobata colonies took longer to bleach, longer to recover and longer to die. In contrast, Acropora hyacinthus and A. millepora colonies bleached quickly and most had either recovered, or died, within 14 wk of the initial reports of bleaching. Whole colony mortality was high in A. hyacinthus (88%) and A. millepora (32%) and partial mortality rare. In contrast, most colonies of P. daedalea and P. lobata lost some tissue and few whole colonies died. The mean proportion of tissue lost per colony was 43±6.6 % and 11±1.1 % respectively. Consequently, observed hierarchies of species susceptibility will depend critically on the time since the onset of stress and must consider both whole and partial colony mortality. Colony mortality was highly dependent on visual estimates of the severity of bleaching but independent of size. Growth rates of Acropora colonies were highly variable and

largely independent of the severity of bleaching. A. hyacinthus was more susceptible to bleaching than A. millepora with 45% of surviving colonies gravid compared to 88%. High whole-colony mortality combined with a reduction in the reproductive output of surviving Acropora suggests that recovery to former levels of abundance is likely to be slow. Bak, RPM and Meesters, EH. (1999) Population structure as a response of coral communities to global change. American Zoologist 39, 56-64. Coral population size structure is generally highly skewed, with a preponderance of the smallest colony size class in populations. To begin to assess possible effects of global change on coral populations, which will be largely controlled in the next decade(s) by "non-climate variables" such as sedimentation, turbidity and nutrient load, we compared degraded reef environments with less degraded reefs. To consider population dynamics within and between parts of coral species metapopulations we use data over two spatial scales (10 and 2,000 km). Colony size distributions appear to be affected in degraded/marginal reefs. This implies changes in mortality patterns (or recruitment) that result in relatively fewer small and more large colonies in populations. We predict that the short-term effects of global change, deterioration of local conditions, will not affect the occurrence of large coral colonies (in terms of absolute size, and possibly mean size) but will limit the abundance of small corals. Long-term global change will increasingly include a component of climate change and the effect on coral populations may become more diverse, although effects such as a decreasing calcium carbonate saturation state will also first affect the abundance of coral recruits. We hypothesize that over the next decade(s) coral populations will become increasingly skewed toward larger colonies. Baker, AC. (2001) Reef corals bleach to survive change. Nature 411, 765-766 No abstract available. Baker, AC, Starger, CJ, McClanahan, TR and Glynn, PW. (2004) Corals’ adaptive response to climate change. Nature 430, 741. Shifting to new algal symbionts may safeguard devastated reefs from extinction. Barton, AD and Casey, KS. (2005) Climatological context for large-scale coral bleaching. Coral Reefs 24, 536-554. Large-scale coral bleaching was first observed in 1979 and has occurred throughout virtually all of the tropics since that time. Severe bleaching may result in the loss of live coral and in a decline of the integrity of the impacted coral reef ecosystem. Despite the extensive scientific research and increased public awareness of coral bleaching, uncertainties remain about the past and future of large-scale coral bleaching. In order to reduce these uncertainties and place large-scale coral bleaching in the longer-term climatological context, specific criteria and methods for using historical sea surface temperature (SST) data to examine coral bleaching-related thermal conditions are proposed by analyzing three, 132 year SST reconstructions: ERSST, HadISST1, and GISST2.3b. These methodologies are applied to case studies at Discovery Bay, Jamaica (77.27°W, 18.45°N), Sombrero

Reef, Florida, USA (81.11°W, 24.63°N), Academy Bay, Galápagos, Ecuador (90.31°W, 0.74°S), Pearl and Hermes Reef, Northwest Hawaiian Islands, USA (175.83°W, 27.83°N), Midway Island, Northwest Hawaiian Islands, USA (177.37°W, 28.25°N), Davies Reef, Australia (147.68°E, 18.83°S), and North Male Atoll, Maldives (73.35°E, 4.70°N). The results of this study show that (1) The historical SST data provide a useful long-term record of thermal conditions in reef ecosystems, giving important insight into the thermal history of coral reefs and (2) While coral bleaching and anomalously warm SSTs have occurred over much of the world in recent decades, case studies in the Caribbean, Northwest Hawaiian Islands, and parts of other regions such as the Great Barrier Reef exhibited SST conditions and cumulative thermal stress prior to 1979 that were comparable to those conditions observed during the strong, frequent coral bleaching events since 1979. This climatological context and knowledge of past environmental conditions in reef ecosystems may foster a better understanding of how coral reefs will respond in future, ocean warming scenarios. Bassim, KM, Sammarco, PW and Snell, TL. (2002) Effects of temperature on success of (self and non-self) fertilization and embryogenesis in Diploria strigosa (Cnidaria, Scleractinia). Marine Biology 140, 479-488. The effects of seawater temperature on the degree of success of self-fertilization versus cross-fertilization and on embryogenesis were investigated in the scleractinian coral Diploria strigosa (Dana 1846). Gametes from nine colonies were collected from the Flower Garden Banks reefs, a set of coral reefs in the northern Gulf of Mexico, ~110 nm offshore from Texas, United States. Gametes from the coral colonies were combined in all possible combinations and placed in 30°, 31°, and 32°C water baths on successive nights, respectively. Each cross, whether selfed or out-crossed, and each temperature treatment, was replicated twice. A high frequency of successful self- and cross-fertilization was observed at all temperature treatments and in both blocks. Higher temperatures, however, commonly produced numerous developmental aberrations during embryogenesis of the larvae. Thus, although fertilization rates can remain high under high temperature conditions, if temperatures remain high for several days, embryonic development and larval viability may be expected to decrease dramatically. We propose that the success of coral larval development may be diminished in areas where abnormally high sea surface temperatures occur during the spawning season. We also propose that highly successful selfing may enhance the abundance of locally adapted genotypes, which in turn may be advantageous where reefs, such as the Flower Garden Banks, are relatively geographically isolated. Bassim, KM and Sammarco, PW. (2003) Effect of temperature and ammonium on larval development and survival in scelractinian coral (Diploria strigosa). Marine Biology 142, 241-252. Increases in ambient water temperature on coral reefs around the world, along with regional nutrient enrichment, have been a growing concern in coral reef ecology. We studied the effects of seawater temperature and ammonium concentrations on the azooxanthellate planular larvae of Diploria strigosa Dana, 1846 (Cnidaria: Scleractinia) over a period of 9 days. We did this to determine whether increases in these environmental variables affect coral larval development and survival.

Settlement frequencies were also examined. Larvae were placed in water baths at 28°C, 30°C, and 32°C (ambient temperature at time of sampling was ~29°C). Larvae in 30°C and 32°C suffered approximately 50% and 70% greater mortality, respectively, than those at 28°C. At each of the three temperatures, separate groups of larvae were exposed to a 20 µM l–1 concentration of NH4

+ (as NH4Cl), a concentration similar to that measured on certain reefs in the Florida Keys. Seawater temperatures of 30–32°C slowed or halted development in the later stages of larval development. At 32°C, time spent by larvae in a swimming/searching mode was observed to be higher than that at 28°C or 30°C. In the 28°C and 30°C treatments, D. strigosa planulae exhibited phototactic responses similar to those of other scleractinian corals–positively phototactic initially and then negatively so after ≥50 h; at seawater temperatures of 32°C, planulae became immediately negatively phototactic. In general, an increase in the seawater temperature caused a significant decrease in ciliary activity (motility) and rate of settlement in the larvae in a manner proportional to temperature. The presence of ammonium also caused a significant decrease in these variables, and these effects were additive with respect to those of increased temperature. The lack of symbiotic algae (which can assimilate ammonia) may have contributed to the observed increased mortality levels under conditions of enriched NH4

+. Calculation of isochrones (distances which a larva may traverse within a given period of time) for planulae exposed to conditions of increased temperature and/or ammonium concentrations suggests that a resultant decrease in larval longevity could potentially decrease distance of larval dispersal. Beck, MW and Odaya, M. (2001) Ecoregional planning in marine environments: Identifying priority sites for conservation in the northern Gulf of Mexico. Aquatic Conservation: Marine and Freshwater Ecosystems 11, 235-242. 1. The overall aim of this work was to identify sites within the northern Gulf of Mexico that, if protected, would fully represent the biological diversity of the nearshore waters of this ecoregion. In this paper, we focus on the eastern subregion, northwest Florida, to illustrate the process of ecoregional planning in marine environments. 2. The basic steps in ecoregional planning include: identification of conservation targets, i.e. species and habitats; collection of data on their ecology and distribution; determination of conservation goals for the amount of targets that must be protected; and identification of a set of sites that meets these goals for all targets. 3. As a preliminary goal, it was determined that the set of priority sites should contain at least 20% of the current distribution of each target habitat and species. 4. Two primary tools were used to choose a set of priority sites: a reserve selection program, SITES, and interviews and a workshop with scientists and managers. The final set of priority sites integrated information from these various sources. 5. The goals were exceeded for all conservation targets in the priority sites. On average, 75 and 58% of the distributions of the habitat targets were contained within the priority and high priority sites, respectively. These priority and high priority sites only occupied 29 and 17% of the study area, respectively. 6. One of the first steps in the conservation of the bays identified as priority sites is the recognition that they are integrally linked by important processes to the surrounding terrestrial and aquatic environments. Conservation in a part of these estuarine landscapes will benefit biodiversity across environments.

Beger, M, Solandt, J-L and Dacles, T. (2001) Coral reef bleaching at Danjugan Island, Negros Occidental, Philippines: A two year monitoring programme. The Philippines Reef and Rainforest Conservation Foundation: Bacolod City (Philippines), 19 pp. Danjugan Island (Negros Occidental, Philippines) is known through anecdotal reports to be a site of high biodiversity. In recent years, partial deforestation for coconut production combined with increased fishing pressure and use of destructive fisheries practices have denuded Danjugan's littoral forest and fringing coral reef systems. This has been a major contributory factor leading to the impoverishment of local, coastal-dwelling communities. In 1994 the Philippine Reef and Rainforest Project was launched to provide biodiversity conservation, restoration and education programmes for Danjugan Island and adjacent coastal areas under a tri-partite collaborative arrangement between the Philippine Reef and Rainforest Conservation Foundation Inc. (PRRCFI), Coral Cay Conservation Ltd (CCC) and the World Land Trust (WLT). Between August and November 1995 and May and July 1996 CCC and PRRCFI volunteers carried out 51 survey transects on the fringing reef around Danjugan Island and two nearby reef complexes to assess their physical and biological characteristics. The gross features of the physical environment, including the dominant winds and currents, were also recorded together with any obvious indications of anthropogenic disturbance. The volunteer training programme and survey techniques used were based on those developed by CCC for similar projects in Belize. Three additional research projects were conducted during this phase of the project: identification of suitable areas as permanent monitoring sites; testing a technique for assessing coral health and testing a simple methodology developed by the International Centre for Living Aquatic Resources Management for surveys of coral reefs. The survey work suggests that the fringing reefs of Danjugan Island are amongst the world's most diverse small reefs, with at least 236 scleractinians and hydrocorals and 229 fish species documented. Of the coral species recorded, 20 are regarded as being rare in the Philippines with possibly one species not previously recorded from the region. Numerous other interesting and unique features were recorded, including four colonies of the rare Acropora monticulosa. Data acquired during this phase of the project were used to create an environmental database for future preparation of habitat maps of the project area. A range of univariate and multivariate techniques were used for data analysis and interpretation of the data This data analysis, combined with anecdotal observations by CCC and PRRCFI scientists, provided a rationale in support of the establishment of management policies for the protection and sustainable use of Danjugan's coastal marine resources. This report documents the results of baseline marine surveys undertaken by CCC and PRRCFI personnel at Danjugan Island in 1995 and 1996. The information within this report, and the four other sister reports has been recently compiled from the large data set that has not been fully analysed until now. The information here will provide the managing team employed by PRRCFI to act on key findings related to the environment inside and outside all three key areas within the Danjugan area; special management areas within the reserve, the reserve itself, and areas outside the area of the reserve.

Bellwood, DR, Hoey, AS and Choat, JH. (2003) Limited functional redundancy in high diversity systems: Resilience and ecosystem function on coral reefs. Ecology Letters 6, 281-285. Biodiversity is frequently associated with functional redundancy. Indo-Pacific coral reefs incorporate some of the most diverse ecosystems on the globe with over 3000 species of fishes recorded from the region. Despite this diversity, we document changes in ecosystem function on coral reefs at regional biogeographical scales as a result of overfishing of just one species, the giant humphead parrotfish (Bolbometopon muricatum). Each parrotfish ingests over 5 tonnes of structural reef carbonates per year, almost half being living corals. On relatively unexploited oceanic reefs, total ingestion rates per m2 balance estimated rates of reef growth. However, human activity and ecosystem disruption are strongly correlated, regardless of local fish biodiversity. The results emphasize the need to consider the functional role of species when formulating management strategies and the potential weakness of the link between biodiversity and ecosystem resilience. Bellwood, DR, Hughes, TP, Folke, C and Nyström, M. (2004) Confronting the coral reef crisis. Nature 429, 827-833. The worldwide decline of coral reefs calls for an urgent reassessment of current management practices. Confronting large-scale crises requires a major scaling-up of management efforts based on an improved understanding of the ecological processes that underlie reef resilience. Managing for improved resilience, incorporating the role of human activity in shaping ecosystems, provides a basis for coping with uncertainty, future changes and ecological surprises. Here we review the ecological roles of critical functional groups (for both corals and reef fishes) that are fundamental to understanding resilience and avoiding phase shifts from coral dominance to less desirable, degraded ecosystems. We identify striking biogeographic differences in the species richness and composition of functional groups, which highlight the vulnerability of Caribbean reef ecosystems. These findings have profound implications for restoration of degraded reefs, management of fisheries, and the focus on marine protected areas and biodiversity hotspots as priorities for conservation. Bellwood, DR, Hoey, AS, Ackerman, JL and Depczynski, M. (2006) Coral bleaching, reef fish community phase shifts and the resilience of coral reefs. Global Change Biology 12, 1587–1594. The 1998 global coral bleaching event was the largest recorded historical disturbance of coral reefs and resulted in extensive habitat loss. Annual censuses of reef fish community structure over a 12-year period spanning the bleaching event revealed a marked phase shift from a prebleach to postbleach assemblage. Surprisingly, we found that the bleaching event had no detectable effect on the abundance, diversity or species richness of a local cryptobenthic reef fish community. Furthermore, there is no evidence of regeneration even after 5–35 generations of these short-lived species. These results have significant implications for our understanding of the response of coral reef ecosystems to global warming and highlight the importance of selecting appropriate criteria for evaluating reef resilience.

Bellwood, DR, Hughes, TP and Hoey, AS. (2006) Sleeping Functional Group Drives Coral-Reef Recovery. Current Biology 16, 2434–2439. The world’s coral reefs are in decline, with many exhibiting a phase shift from coral to macroalgal dominance. This change is often associated with habitat loss and overharvesting of herbivorous fishes, particularly parrotfishes and surgeonfishes. The challenge is to reverse this decline and enhance the resilience of coral-reef ecosystems.We demonstrate, by using a large-scale experimentally induced phase shift, that the rapid reversal from a macroalgal-dominated to a coral- and epilithic algal-dominated state was not a result of herbivory by parrotfishes or surgeonfishes. Surprisingly, phase-shift reversal was primarily driven by a single batfish species (Platax pinnatus), a fish previously regarded as an invertebrate feeder. The 43 herbivorous fishes in the local fauna played only a minor role, suggesting that biodiversity may not offer the protection we hoped for in complex ecosystems. Our findings highlight the dangers faced by coral reefs and other threatened complex ecosystems: Species or functional groups that prevent phase shifts may not be able to reverse phase shifts once they occur. Nevertheless, reversal is possible. The critical issue is to identify and protect those groups that underpin the resilience and regeneration of complex ecosystems. Bena, C and van Woesik, R. (2004) The impact of two bleaching events on the survival of small coral colonies (Okinawa, Japan). Bulletin of Marine Science 75, 115-125. This study addresses the question whether the density and composition of small coral colonies (≤ 3 cm diameter) were affected by the high sea surface temperature anomalies in the summers of 1998 and 2001 in western Okinawa, Japan. While the percentage of coral cover decreased dramatically at depths < 4 m in 1998, there were more subtle changes in the abundance of small coral colonies. Change in the latter was attributed mainly to mortality of Acropora, Montipora, and pocilloporid colonies on shallow reefs, although the populations recovered within 2 yrs because of recruitment. Small coral colonies were less affected by the 2001 thermal anomaly than by the 1998 event, although they were of similar intensity and duration. The density of small Porites colonies changed little in 1998, but showed a steady decline after 1998, while there was no obvious change in the density of faviids. The limited impact of elevated temperatures and high irradiance on small coral colonies and the significant impact on large colonies concur with mass transfer theory, which suggests that small colonies have higher mass transfer rates than large colonies and are hence more likely to survive in times of heat and irradiance stress. Bengtsson, J, Angelstam, P, Elmqvist, T, Emanuelsson, U, Folke, C, Ihse, M, Moberg, F and Nyström, M. (2002) Reserves, resilience and dynamic landscapes. Ambio 32, 389-396. In a world increasingly modified by human activities, the conservation of biodiversity is essential as insurance to maintain resilient ecosystems and ensure a sustainable flow of ecosystem goods and services to society. However, existing reserves and national parks are unlikely to incorporate the long-term and large-scale dynamics of ecosystems. Hence, conservation strategies have to actively incorporate the large

areas of land that are managed for human use. For ecosystems to reorganize after large-scale natural and human-induced disturbances, spatial resilience in the form of ecological memory is a prerequisite. The ecological memory is composed of the species, interactions and structures that make ecosystem reorganization possible, and its components may be found within disturbed patches as well in the surrounding landscape. Present static reserves should be complemented with dynamic reserves, such as ecological fallows and dynamic successional reserves, that are part of ecosystem management mimicking natural disturbance regimes at the landscape level. Berkelmans, R and Willis, BL. (1999) Seasonal and local spatial patterns in the upper thermal limits of corals on the inshore Central Great Barrier Reef. Coral Reefs 18, 219-228. Experimental studies of the upper thermal limits of corals from Orpheus Island, an inshore reef in the central Great Barrier Reef, show that Acropora formosa has a 5-day 50%-bleaching threshold of between 31 and 32°C in summer, only 2 to 3°C higher than local mean summer temperatures (29°C). Summer bleaching thresholds for Pocillopora damicornis and A. elseyi were 1°C higher (between 32 and 33°C). The winter bleaching threshold of Pocillopora damicornis was 1°C lower than its summer threshold, indicating that seasonal acclimatisation may take place. This seasonal difference raises the possibility that at least some corals may be capable of short-term thermal acclimatisation. Neither P. damicornis nor A. elseyi showed habitat-specific (reef flat versus reef slope) differences in bleaching thresholds. Further, colonies of P. damicornis collected from sites 3 km apart also showed no difference in bleaching threshold despite populations of this species responding differently at these two sites during a natural bleaching event. The bleaching thresholds determined in this study are best considered as the maximum tolerable temperatures for local populations of these species because they were determined in the absence of additional stressors (e.g. high light) which often co-occur during natural bleaching events. We consider the 5-day 50% bleaching thresholds determined in these experiments to be fair indicators of upper thermal limits, because >50% of a sample population died when allowed to recover in situ. We found a delay of up to a month in the bleaching response of corals following thermal stress, a result that has implications for identifying the timing of stressful conditions in natural bleaching events. Berkelmans, R. (2002) Time-integrated thermal bleaching thresholds of reefs and their variation on the Great Barrier Reef. Marine Ecology Progress Series 229, 73-82. Knowledge of the critical levels for key environmental variables that are likely to cause bleaching in reef corals is of fundamental importance in conducting risk assessments of potential climate-change effects on coral reefs. Such knowledge can also be used to provide early warning of mass bleaching events. A number of factors have contributed to the difficulty in determining critical levels for coral bleaching. These factors include the fact that multiple stressors may be involved in bleaching, the duration of stress required to elicit a bleaching response varies with temperature, and bleaching triggers are known to be variable in space, time and by species. In this study, I identify sea surface temperature (SST) as the most important parameter for

predicting coral bleaching from 4 possible environmental variables collected over 10 to 12 yr from weather stations at 2 locations on the Great Barrier Reef (GBR): temperature, wind speed, solar radiation and barometric pressure. Predicted bleaching-response curves are constructed from high-resolution in situ temperature records and historical observations of coral bleaching for 13 locations. These curves approximate reef-wide stress-response thresholds for bleaching of thermally sensitive (and often dominant) coral species. Distinct spatial trends exist in the thermal sensitivity of coral populations that correspond with position across the shelf and latitude in the case of mid- and outer-shelf reefs. This suggests that considerable thermal adaptation has taken place over small (10s of km) and large (100s to 1000s of km) spatial scales. Bleaching curves for inshore reefs do not correspond with latitude and are more variable, reflecting greater local-scale variability in temperature regimes. Berkelmans, R, Done, T, Goggin, L and Harriott, V. (2002) Coral bleaching and global climate change: Current state of knowledge. CRC Reef Research Centre: Townsville (Australia), 6 pp. Coral bleaching is the loss of colour from corals under stressful environmental conditions. While any stress can cause corals to bleach, high water temperature has been the major cause of coral bleaching events worldwide in recent decades. In 1998, a major bleaching event in inshore reef waters of the Great Barrier Reef raised concerns about the health of the reef. With predictions that temperatures will continue to rise as a result of global warming, the future of tropical coral reefs is causing concern worldwide. Berkelmans, R, De’ath, G, Kininmonth, S and Skirving, WJ. (2004) A comparison of the 1998 and 2002 coral bleaching events on the Great Barrier Reef: Spatial correlation, patterns, and predictions. Coral Reefs 23, 74-83. Detailed mapping of coral bleaching events provides an opportunity to examine spatial patterns in bleaching over scales of 10’s to 1,000’s of km and the spatial correlation between sea surface temperature (SST) and bleaching. We present data for two large-scale (2,000 km) bleaching events on the Great Barrier Reef (GBR): one from 1998 and another from 2002, both mapped by aerial survey methods. We examined a wide range of satellite-derived SST variables to determine which one best correlated with the observed bleaching patterns. We found that the maximum SST occurring over any 3-day period (max3d) during the bleaching season predicted bleaching better than anomaly-based SST variables and that short averaging periods (3–6 days) predicted bleaching better than longer averaging periods. Short periods of high temperature are therefore highly stressful to corals and result in highly predictable bleaching patterns. Max3d SST predicted the presence/absence of bleaching with an accuracy of 73.2%. Largescale (GBR-wide) spatial patterns of bleaching were similar between 1998 and 2002 with more inshore reefs bleached compared to offshore reefs. Spatial change in patterns of bleaching occurred at scales of ~10’s km, indicating that reefs bleach (or not) in spatial clusters, possibly due to local weather patterns, oceanographic conditions, or both. Approximately 42% of reefs bleached to some extent in 1998 with ~18% strongly bleached, while in 2002, ~54% of reefs bleached to some extent with ~18% strongly bleached. These

statistics and the fact that nearly twice as many offshore reefs bleached in 2002 compared to 1998 (41 vs. 21%, respectively) makes the 2002 event the worst bleaching event on record for the GBR. Modeling of the relationship between bleaching and max3d SST indicates that a 1°C increase would increase the bleaching occurrence of reefs from 50% (approximate occurrence in 1998 and 2002) to 82%, while a 2°C increase would increase the occurrence to 97% and a 3°C increase to 100%. These results suggest that coral reefs are profoundly sensitive to even modest increases in temperature and, in the absence of acclimatization/adaptation, are likely to suffer large declines under mid-range International Panel for Climate Change predictions by 2050. Berkelmans, R and van Oppen, MJ. (2006) The role of zooxanthellae in the thermal tolerance of corals: A ‘nugget of hope’ for coral reefs in an era of climate change. Proceedings of the Royal Society of London B Biological Sciences 273, 2305-2312. The ability of coral reefs to survive the projected increases in temperature due to global warming will depend largely on the ability of corals to adapt or acclimatize to increased temperature extremes over the next few decades. Many coral species are highly sensitive to temperature stress and the number of stress (bleaching) episodes has increased in recent decades. We investigated the acclimatization potential of Acropora millepora, a common and widespread Indo-Pacific hard coral species, through transplantation and experimental manipulation. We show that adult corals, at least in some circumstances, are capable of acquiring increased thermal tolerance and that the increased tolerance is a direct result of a change in the symbiont type dominating their tissues from Symbiodinium type C to D. Our data suggest that the change in symbiont type in our experiment was due to a shuffling of existing types already present in coral tissues, not through exogenous uptake from the environment. The level of increased tolerance gained by the corals changing their dominant symbiont type to D (the most thermally resistant type known) is around 1-1.5 degrees C. This is the first study to show that thermal acclimatization is causally related to symbiont type and provides new insight into the ecological advantage of corals harbouring mixed algal populations. While this increase is of huge ecological significance for many coral species, in the absence of other mechanisms of thermal acclimatization/adaptation, it may not be sufficient to survive climate change under predicted sea surface temperature scenarios over the next 100 years. However, it may be enough to 'buy time' while greenhouse reduction measures are put in place. Berumen, ML and Pratchett, MS. (2006) Recovery without resilience: persistent disturbance and long-term shifts in the structure of fish and coral communities at Tiahura Reef, Moorea. Coral Reefs 25, 647–653. Disturbances have a critical effect on the structure of natural communities. In this study long-term changes were examined in the reef community at Tiahura Reef, on the northern coast of Moorea, which had been subject to many and varied disturbances over the last 25 years. Tiahura Reef was subject to an outbreak of crown-of-thorns starWsh (Acanthaster planci) in 1980–1981, causing significant declines in the abundance of scleractinian corals and butterXyWshes. By 2003, the abundance of corals and butterflyfishes had returned to former levels, but despite this apparent recovery, the species composition of coral communities and butterflyfish

assemblages was very different from those recorded in 1979. Ongoing disturbances (including further outbreaks of crown-of-thorns starWsh, cyclones, and coral bleaching events) appear to have prevented recovery of many important coral species (notably, Acropora spp.), which has had subsequent effects on the community structure of coral-feeding butterflyfishes. This study shows that recurrent disturbances may have persistent effects on the structure and dynamics of natural communities. Best, BA, Pomeroy, RS and Balboa, CM. (2002) Implications for coral reef management and policy: Relevant findings from the 9th International Coral Reef Symposium. 9th International Coral Reef Symposium: Bali (Indonesia), 113 pp. Coral reefs constitute one of the earth’s most complex, beautiful and biologically diverse ecosystems. These unique ecosystems benefit people directly by supplying a vast array of goods and services such as food, medicine, recreation, and coastal protection, as well as aesthetic and cultural benefits. Coral reefs occur in over 100 countries. According to one estimate, coral reefs provide over US $375 billion worth of goods and ecosystem services to humans. The economies of many countries, especially small island nations, are highly dependent on the goods and services that coral reefs provide. In addition, coral reefs are intimately associated with mangrove forests and seagrass meadows and form a broader tropical coastal ecosystem upon which more than a billion people depend. Unfortunately, many coral reefs around the world are in serious decline. According to the recent report Status of Coral Reefs of the World: 2000 by the Global Coral Reef Monitoring Network, one quarter of the world’s reefs have already been lost and another one-third may disappear within the next 30 years. Coral reefs are threatened both directly and indirectly from a variety of human activities. These threats include coastal development, overexploitation and destructive fishing practices, diseases, land-based pollution and erosion, marine-based pollution, and global climate change. In addition, the recent global impacts of catastrophic events, such as widespread coral bleaching and mortality and increased storm intensity, compound the more localized human impacts that place reefs at risk. There is an urgent need to respond to these threats facing coral reefs at local, national, regional, and global levels in order to address biodiversity loss, food insecurity, loss of economic livelihood, and loss of development potential. In response to these threats facing coral reefs, the organizers of the 9th International Coral Reef Symposium (ICRS) incorporated strong management and human or socio-economic dimensions into the symposium, expanding their usual biological and ecological emphasis. Over 1500 scientists, managers, resource users, government officials, journalists and others interested in coral reef studies and management gathered in Bali, Indonesia in October of 2000. The overall theme of the meeting was the World Coral Reefs in the New Millennium: Bridging Research and Management for Sustainable Development. To intervene effectively, we need to view and understand coral reefs in multiple dimensions – human, socioeconomic, biological, and ecological. The 9th ICRS continues a process begun in 1969 at the first ICRS – bringing together those interested in coral reef studies and management to share, debate and learn from each other, and to set a course of action to conserve and sustain the coral reefs. Organized into five broad themes: “State of Knowledge;” “Resource Management;” “Socioeconomic Values;” “Assessment, Monitoring, and

Rehabilitation;” and “The Future of Coral Reefs,” over 1400 papers were presented orally in 58 mini-symposia, and more papers were presented in poster sessions. The purpose of this report is to synthesize some of the best scientific and management information presented at ICRS for use by those in positions to conserve, protect and rehabilitate coral reefs – policy-makers, managers and the public-at-large. Session convenors, collaborators and colleagues were asked to prepare short syntheses of relevant sessions and/or topics for coral reef management and policy. Each topic area is meant to be a stand-alone piece that can be used separately from the whole report. Some topics relate directly to one or more sessions, while other topics were dispersed throughout many sessions. We extend our sincere thanks and gratitude to Dr. Anugerah Nontji and his Indonesian colleagues for hosting a wonderful Symposium, and to those session convenors and colleagues who responded to our requests for contributions to this report. The success of this project is due to their dedication and assistance. Although efforts must be made by all parties on all issues, some of the more important recommendations from the 9th ICRS are the following: For Policymakers � reduce greenhouse gases and address climate change � address threats from invasive species and coral diseases � strengthen law enforcement � reduce land-based sources of marine pollution � expand and strengthen marine protected areas � enhance communication of scientific information to the general public For Researchers � encourage research targeted to management needs � conduct valuation studies of coral reefs � increase monitoring studies for more informed management For Managers � implement co-management approaches � implement ecosystem level management � make more use of socioeconomic information in management � promote stakeholder participation and participatory decision-making in management � exercise vigilance and precautionary approaches in all coral reef fisheries � prohibit destructive fishing practices – such as explosives, cyanide and other poisons, dredging, and trawling We must now utilize the information available to us to conserve and protect the world’s coral reefs, in conjunction with a precautionary approach to management. We must also develop policy and management actions that reflect the multiple dimensions of coral reef ecosystems – human, biological and ecological. Bhagooli, R and Hidaka, M. (2003) Comparison of stress susceptibility of in hospite and isolated zooxanthellae among five coral species. Journal of Experimental Marine Biology and Ecology 291, 181-197. Coral species in a similar habitat often show different bleaching susceptibilities. It is not understood which partner of coral–zooxanthellae complexes is responsible for differential stress susceptibility. Stress susceptibilities of in hospite and isolated zooxanthellae from five species of corals collected from shallow water in Okinawa were compared. To estimate stress susceptibility,we measured the maximum

quantum yields (Fv/Fm) of in hospite and isolated zooxanthellae after 3-h exposure to either 28 or 34°C at various light intensities and their recovery after 12 h under dim light at 26°C. Significant reduction in photochemical efficiency (Fv/Fm) of photosystem II (PSII) was observed in in hospite zooxanthellae exposed to high light intensity (1000 µmol quanta m-2 s-1), while PSII activity of isolated zooxanthellae decreased significantly even at a lower light intensity (70 µmol quanta m-2 s-1). The recovery of the PSII activity after 12 h was incomplete in both in hospite and isolated zooxanthellae, indicating the presence of chronic photoinhibition. The stress susceptibility of isolated zooxanthellae was more variable among species than in hospite zooxanthellae. The order of stress susceptibility among the five coral species was different between in hospite and isolated zooxanthellae. The present results suggest that the host plays a significant role in determining bleaching susceptibility of corals, though zooxanthellae from different host have different stress susceptibilities. Bhagooli, R and Hidaka, M. (2004) Photoinhibition, bleaching susceptibility and mortality in two scleractinian corals, Platygyra ryukyuensis and Stylophora pistillata in response to thermal and light stresses. Comparative Biochemistry and Physiology. Part A, Molecular and Integrative Physiology 137, 547-555. In the present study, we examined the effect of thermal stress on the photoinhibitory light threshold in a bleaching susceptible (Stylophora pistillata) and a bleaching resistant (Platygyra ryukyuensis) coral. Four light (0, 110, 520, 1015 micromol quantam(-2)s(-1)) and three temperature (26, 32 and 34 degrees C) conditions were used over a 3-h period, followed by 24- and 48-h recovery periods at approximately 21 degrees C under dim light. Dynamic photoinhibition could be detected in both P. ryukyuensis and S. pistillata under 520 and 1015 micromol quantam(-2)s(-1) at 26 degrees C and under 110 micromol quantam(-2)s(-1) at 32 degrees C only in S. pistillata. Chronic photoinhibition was recorded under 520 and 1015 micromol quantam(-2)s(-1) at 34 degrees C in P. ryukyuensis, and under 1015 micromol quantam(-2)s(-1) at 32 degrees C and under all light levels at 34 degrees C in S. pistillata. These results show that high temperature reduced the threshold light intensity for photoinhibition differently in two corals with different bleaching susceptibilities under thermal stress. No visual paling and mortality in P. ryukyuensis was observed at any treatment, even in chronically photoinhibited specimens, while paling and high mortality of S. pistillata was noted in all treatments, apart from samples at 26 degrees C. These observations suggest a potential role of the host in differential bleaching and mortality determination. Bhagooli, R and Hidaka, M. (2004) Release of zooxanthellae with intact photosynthetic activity by the coral Galaxea fascicularis in response to high temperature stress. Marine Biology 145, 329-337. Damage to zooxanthellae photosynthetic apparatus has been proposed to be the underlying mechanism of coral bleaching, but how the expulsion of zooxanthellae is triggered is still not understood. The present study compared the photosystem II (PSII) functioning and overall photosynthesis of retained and released zooxanthellae from the reef-building coral Galaxea fascicularis exposed to high temperature stress. The use of pulse-amplitude-modulated (PAM) fluorometry for isolated zooxanthellae was validated and used to measure photosynthesis. There was no significant difference in PSII functioning and the overall photosynthesis between retained

zooxanthellae, which were isolated immediately after stress treatment, and those released from the coral exposed to either 30 or 32°C, while the zooxanthellae population released at 28°C showed significantly lower PSII functioning than those retained in the polyps. The release of healthylooking zooxanthellae by polyps exposed to elevated temperatures was significantly higher than those in the control (28°C). Higher release of undischarged cnidae, indicative of host cell necrosis or detachment, was observed in 32°C treatments. These findings indicate that the zooxanthellae released in 30 or 32°C treatments exhibited normal morphology and intact photosynthetic activity. The present results strongly suggest that the release of zooxanthellae from G. fascicularis at 30 or 32°C is a non-selective process with respect to the zooxanthellar PSII functioning and thus the host seems to be the first partner to be physiologically affected in temperature-induced bleaching. Birrell, CL, McCook, LJ and Willis, BL. (2005) Effects of algal turfs and sediment on coral settlement. Marine Pollution Bulletin 51, 408-414. Successful settlement and recruitment of corals is critical to the resilience of coral reefs. Given that many degraded reefs are dominated by benthic algae, recovery of coral populations after bleaching and other disturbances requires successful settlement amidst benthic algae. Algal turfs often accumulate sediments, sediments are known to inhibit coral settlement, and reefs with high inputs of terrestrial sediments are often dominated by turfs. We investigated the impacts of two algal turf assemblages, and of sediment deposits, on settlement of the coral Acropora millepora (Ehrenberg). Adding sediment reduced coral settlement, but the effects of different algal turfs varied. In one case, algal turfs inhibited coral settlement, whereas the other turf only inhibited settlement when combined with sediments. These results provide the first direct, experimental evidence of effects of filamentous algal turfs on coral settlement, the variability in those effects, and the potential combined effects of algal turfs and trapped sediments. Booth, DJ and Beretta, GA. (2002) Changes in fish assemblage after a coral bleaching event. Marine Ecology Progress Series 245, 205-212. Large-scale bleaching events are major disturbances to coral health and community structure, but may also affect other coral reef taxa, such as fishes. In 1997-1998, wide-spread coral bleaching and mortality occurred across the Great Barrier Reef, caused by increases in sea surface temperatures resulting from El Niño Southern Oscillation. As part of this event, in February-March 1998, there was extensive bleaching within One Tree Island lagoon (southern Great Barrier Reef), such that almost 12 mo later there was a significant reduction of live coral cover at some sites. We monitored the distribution of adults and recruitment of damselfishes (Pomacentridae) to sites within One Tree Island in 1993-1995 and in 1999. Fish species that normally associate with live corals showed relatively lower recruitment at bleached sites compared to the same sites pre-bleaching or to recovered sites and, as a result, species diversity and assemblage structure of recruits changed. Compared to 1993/1995 densities, adult Pomacentrus wardi, P. moluccensis and Chrysiptera rollandi densities dropped at bleached sites, but not at unbleached sites. While P. moluccensis directly associate with live corals, the other 2 species do not. This study has demonstrated that indirect effects of bleaching can include changes in assemblage structure of reef fish adults and recruits.

Briggs, JC. (2005) Coral reefs: Conserving the evolutionary sources. Biological Conservation 126, 297-305. The current worldwide degradation of coral reefs constitutes an international problem that calls for immediate attention. A multitude of conservation hotspots scattered over the circumtropical seas have been identified, but there has been no general agreement as to how to attack the problem. The major difficulty seems to be the lack of a priority system. Are some reefs more important than others? If so, where are they located and what needs to be done? Some reefs are located in areas where the biodiversity (species, genera, families) is exceptionally high and where there is evidence of past and present evolutionary production. These areas are called centers of origin and one is located in the Indo-Pacific Ocean and the other in the Atlantic. The Indo-Pacific centre, located in the East Indies Triangle, is considered to be the primary tropical center due to its broad geographic influence. The Atlantic centre, located in the southern Caribbean Sea, is considered a secondary centre because its influence is restricted to the tropical Atlantic. These centres, although occupying relatively small geographic areas, are evidently the source of dominant species that have played the major role in assembling coral reef communities across each ocean. The existence of the two centres permits one to realize that, in the coral reef environment, there is a geographical separation between process and pattern. Recognition of the separation will allow significant improvements in conservation planning. The evolutionary process in the centres must be protected in order to maintain the outlying diversity patterns. Brown, BE. (1997) Coral bleaching: Causes and consequences. Coral Reefs 16, S129-S138. It has been over 10 years since the phenomenon of extensive coral bleaching was first described. In most cases bleaching has been attributed to elevated temperature, but other instances involving high solar irradiance, and sometimes disease, have also been documented. It is timely, in view of our concern about worldwide reef condition, to review knowledge of physical and biological factors involved in bleaching, the mechanisms of zooxanthellae and pigment loss, and the ecological consequences for coral communities. Here we evaluate recently acquired data on temperature and irradiance-induced bleaching, including long-term data sets which suggest that repeated bleaching events may be the consequence of a steadily rising background sea temperature that will in the future expose corals to an increasingly hostile environment. Cellular mechanisms of bleaching involve a variety of processes that include the degeneration of zooxanthellae in situ, release of zooxanthellae from mesenterial filaments and release of algae within host cells which become detached from the endoderm. Photo-protective defences (particularly carotenoid pigments) in zooxanthellae are likely to play an important role in limiting the bleaching response which is probably elicited by a combination of elevated temperature and irradiance in the field. The ability of corals to respond adaptively to recurrent bleaching episodes is not known, but preliminary evidence suggests that phenotypic responses of both corals and zooxanthellae may be significant.

Brown, BE, Ambarsari, I, Warner, ME, Fitt, WK, Dunne, RP, Gibb, SW and Cummings, DG. (1999) Diurnal changes in photochemical efficiency and xanthophyll concentrations in shallow water reef corals: evidence for photoinhibition and photoprotection. Coral Reefs 18, 99-105. Diurnal patterns of photoinhibition have been identified in seven species of shallow water reef corals from the Andaman Sea, o! the west coast of Thailand, using pulse amplitude fluorometry. Photochemical efficiency (Fv/Fm) and quantum yield (∆F/ Fm’) of symbiotic dinoflagellates within the corals declined after dawn to reach a minimum between midday and early afternoon, recovering to former dawn levels by early evening. Parallel studies on the xanthophylls diadinoxanthin (Dn) and diatoxanthin (Dt), and their inter-conversion, also revealed a strong diurnal pattern as well as inverse correlations between the xanthophyll ratio Dt/(Dn+Dt) and Fv/Fm and ∆F/ Fm’. These findings suggest a photoprotective function for these pigments. Brown, BE, Dunne, RP, Ambarsari, I, Le Tissier, MDA and Satapoomin, U. (1999) Seasonal fluctuations in environmental factors and variations in symbiotic algae and chlorophyll pigments in four Indo-Pacific coral species. Marine Ecology Progress Series 191, 53-69. Marked seasonality is observed in sea-surface temperature (SST) and photosynthetically active radiation (PAR) received by shallow water corals at Phuket, Thailand (7" N latitude). This seasonality is reflected in both algal densities and algal chlorophylls (a and C2) in 4 massive species of scleractinian corals, namely Coeloseris rnayeri, Goniastrea retiformis, Porites lutea, and G. aspera. Algal numbers and algal chlorophylls are generally maximal at the end of the wet season (November) and minimal at the end of the dry season (March to May). Algal densities are reduced by up to 53 % in May compared to November in some species, whde algal chlorophylls may be up to 4-fold greater in November. Concomitant with these changes are differences in algal cell volume in 3 out of the 4 species studied and alterations in host tissue depth, with smaller algal volume and greater tissue depth at the end of the wet compared with the dry season. Over the period 1993 to 1996 algal numbers showed a significant negative correlation with monthly (30 d) and short-term (1 to 2 d) values of both SST and PAR dose, while algal chlorophylls displayed a significant negative correlation with PAR dose in the 5 d preceding collection. Rising SST and increasing PAR in the dry season are paralleled by decreasing algal numbers and algal chlorophylls on an annual basis. Recovery of algal densities and a build-up in algal chlorophylls follow reductions in SST and PAR at the end of the wet season. In one of the years, when coral 'bleaching' was widespread at the study site (May 1995), the interaction between SST and PAR resulted in an 86 % reduction in algal densities in bleached corals compared to their annual maximum density. This study highlights the importance of seasonal fluctuations in physical environmental parameters and concomitant changes in the coravalgal symbiosis. Furthermore, it emphasises the need for greater understanding of the potential effects of seasonal and inter-annual variability on the status of algal symbionts, as well as identification of critical times of the year when visible bleaching may not be obvious but when algal numbers and algal chlorophylls are low and effects on coral physiology may be far-reaching.

Brown, BE, Dunne, RP, Goodson, MS and Douglas, AE. (2000) Bleaching patterns in reef corals. Nature 404, 142-143. Coral reefs are under threat from the effects of bleaching, in which symbiotic algae or their photosynthetic pigments are destroyed by increased sea temperatures and solar radiation. Here we show that the bleaching susceptibility of Goniastrea aspera, a shallow-water Indo-Pacific coral, can be predicted from its history of exposure to solar radiation, demonstrating how experience can shape coral bleaching patterns. Brown, BE, Clarke, KR and Warwick, RM. (2002) Serial patterns of biodiversity change in corals across shallow reef flats in Ko Phuket, Thailand, due to effects of local (sedimentation) and regional (climatic) perturbations. Marine Biology 141, 21-29. A 17-year monitoring programme of reef flats at Ko Phuket, Thailand afforded an opportunity to evaluate both univariate and multivariate measures of environmental stress in an assessment of change on coral reef ecosystems. The sites at Ko Phuket suffered the effects of dredging in 1986–1987 and then anomalously low sea levels in 1997–1998 as a result of climate-related events in the Indian Ocean. Univariate measures of species diversity and taxonomic distinctness (D*) increased across the reef flat, reflecting the greater effects of physical stresses on the inner reef flats (compared with outer reef flats) at all sites, with more congeneric species present on the inner reef and more confamilial species on the outer reef. Multivariate measures showed a clear breakdown in seriation patterns at all sites during the dredging in 1987 and in 1998, as a result of earlier negative sea-level anomalies. Recovery from environmental disturbances was obvious within 12 months in each case. The domination of the reefs by massive coral species, which are physiologically adapted to intertidal living and which display partial rather than total colony mortality, may be a factor contributing to the apparent resilience of the reef flats together with continued recruitment and survival of juvenile corals during adverse environmental conditions. Elevated sea temperatures and extensive bleaching of corals in 1991, 1995, and 1998 had no effect upon coral community measures, with many corals recovering their zooxanthellae numbers within 3–5 months of the bleaching events. Brown, BE, Downs, CA, Dunne, RP and Gibb, SW. (2002) Exploring the basis of thermotolerance in the reef coral Goniastrea aspera. Marine Ecology Progress Series 242, 119-129. The shallow-water reef coral Goniastrea aspera Verill 1865 has previously been reported to demonstrate differences in within-colony susceptibility to bleaching at elevated sea temperatures; parts of the colony which are exposed to the highest solar radiation are more thermotolerant than areas which are less exposed. In this paper, we show that at elevated experimental sea temperature the ‘high light’ surfaces lose fewer symbiotic algae, have lower levels of oxidative stress, higher levels of host antioxidant-enzyme copper zinc superoxidase dismutase (CuZnSOD), and host heat-shock proteins 60 and 70, compared to the less exposed surfaces. In addition, ‘high light’ surfaces show less chronic photoinhibition and greater Photosystem II (PS II) recovery potential when exposed to high irradiance at ambient sea temperature. In contrast, no differences were noted in algal defences (e.g. antioxidant enzymes and stress protein production, and xanthophyll cycling) either at

elevated or ambient temperatures. These results are noteworthy because they suggest that corals which acclimatise to high irradiance can, as a result, develop increased thermotolerance which my prevent bleaching at high sea temperatures. importantly, they also demonstrate the significance f the host tissues in maintaining the intact symbiosis of G. aspera under thermal stress. Brown, BE, Dunne, RP, Goodson, MS and Douglas, AE. (2002) Experience shapes the susceptibility of a reef coral to bleaching. Coral Reefs 21, 119-126. Individual zooxanthellate corals vary in their susceptibility to bleaching, caused by the loss of symbiotic algae in response to increased temperatures and solar radiation. In 1995 at Phuket, Thailand, the west face of colonies of the massive coral Goniastrea aspera displayed bleaching in January–March, mediated by high solar radiation. By May, when sea temperatures were anomalously high but solar radiation much reduced, the east sides preferentially bleached. Cores from the east faces of corals exposed to elevated temperature under low irradiance in a laboratory experiment also bleached in response to elevated temperature. The algae in G. aspera were Symbiodinium sp., and the rDNA (both ssu and ITS regions) sequences did not differ between the east and west faces, indicating that the bleaching responses cannot be attributed to genetic factors in the symbiotic algae of the type described in other coral species. It is concluded that, in this system, bleaching is shaped by experience, and that the algae may acquire cross-protection against bleaching from solar to temperature stresses. The impact of experience on bleaching susceptibility may become increasingly important in shaping coral responses to temperature and solar radiation stresses in the coming decades. Bruno, JF, Siddon, CE, Witman, JD, Colin, PL and Toscano, MA. (2001) El Niño related coral bleaching in Palau, Western Caroline Islands. Coral Reefs 20, 127-136. Mass coral bleaching is currently viewed as a major threat to the long-term health of coral reef communities. Here we quantify coral bleaching in Palau coincident with the 1997/1998 El Niño Southern Oscillation event and with local sea surface temperatures of 31 °C, which were 1.0-1.25 °C higher than long-term, satellite derived climatological maximum monthly means for the region. We sampled nine sites, including protected lagoon and fringing reefs, vertical reef walls, and exposed barrier reefs. The percentage of living scleractinian coral tissue that was bleached was 53.4 ± 6.2 (range: 32.3-79.3, n = 8 sites) at 3-5 m depth and 68.9 ± 6.2 (45.7 91.7, n=6 sites) at 10-12 m and did not differ significantly between depths. The overall mean percent cover of bleached scleractinians was 18.9 ± 1.5 (mean ± 1 SE, n = 9 sites), while the cover of healthy corals was only 15.6 ± 2.0. Nearly half (48%) of 946 surveyed colonies belonging to 20 scleractinian taxa were totally bleached, while 15% were partially bleached. Overall, the results indicate that the 1998 coral bleaching episode in Palau was relatively severe and widespread across depths, sites, habitats, and coral taxa. Buddemeier, RW and Fautin, DG. (1993) Coral bleaching as an adaptive mechanism: A testable hypothesis. BioScience 5, 320-326. We posit that bleaching allows a host to be repopulated with a different partner.

Buddemeier, RW, Kleypas, JA and Aronson, RB. (2004) Coral reefs and global climate change: Potential contributions of climate change to stresses on coral reef ecosystems. Pew Center on Global Climate Change: Arlington (USA), 44 pp. Coral reefs are striking, complex, and important features of the marine environment. Reefs are geologic formations constructed from the accumulated skeletons of limestone-secreting animals and plants. The intimately linked plant-animal communities that create them are representative of an ecosystem that occurs in tropical and subtropical waters across the planet, most commonly in shallow oceanic water, and often close to land. Coral reefs have the highest biodiversity of any marine ecosystem, and they provide important ecosystem services and direct economic benefits to the large and growing human populations in low-latitude coastal zones. The natural habitat of coral reefs near the junction of land, sea, and air is both varied and variable, and is a potentially stressful environment. Reef organisms have evolved adaptations over hundreds of millions of years to cope with recurring disturbances: damage or destruction, followed by recovery or regrowth. These are natural features of coral reef history. However, recent global increases in reef ecosystem degradation and mortality (the “coral reef crisis”) appear to be sending a clear message that the rate and nature of recent environmental changes are frequently exceeding the adaptive capacity of coral reef organisms and communities. The coral reef crisis is almost certainly the result of complex and synergistic interactions among local-scale human-imposed stresses and global-scale climatic stresses. Both can produce direct and indirect chronic and acute stresses, leaving few, if any, parts of the ocean truly hospitable for healthy coral reef communities. Documented human stresses include increased nutrient and sediment loading, direct destruction, coastal habitat modification, contamination, and the very important chronic indirect effects of overfishing. The major climate change factor that is becoming increasingly important for coral reefs is rising ocean temperatures, which have been implicated in chronic stress and disease epidemics, as well as in the occurrence of mass coral bleaching episodes. Also of concern are the effects of increasing atmospheric carbon dioxide (CO2) on ocean chemistry, which can inhibit calcification—the deposition of the calcium carbonate minerals that are the structural building materials of coral reefs. Coral reef communities usually recover from acute physical damage or coral mortality if chronic environmental stresses (such as reduced water quality) are weak, and if the acute stresses are not strong or overly frequent. Coral reefs also withstand chronic stresses in the absence of acute stresses. The combination of acute and chronic stress, however, often results in the replacement of the coral reef community by seaweeds or some other nonreef system. Such ecosystem shifts are well advanced in the Caribbean region, where two of the major reef-building coral species have been devastated by disease. In the Indo-Pacific region, the repeated and lethal episodes of “bleaching” associated with unusually high water temperature raise concern that reefs cannot sufficiently recover between such events. Whereas remote oceanic reefs will be affected primarily by climate change, reefs close to human populations will continue to be affected by combinations of additional stresses (e.g., reduced water quality, physical damage, and overharvesting) that must be considered together to be understood and managed. Predictions of the future of coral reefs are difficult because current environmental changes are leading to a combination of surface ocean chemistry and temperature conditions that have

almost certainly never occurred over the evolutionary history of modern coral reef systems. This report reviews the published literature in an effort to analyze the current state of knowledge regarding coral reef communities and the potential contribution of future climate change to coral reef degradation and loss. The major conclusions of the review are summarized as follows: 1) Climate and localized nonclimate stresses interact, often synergistically, to affect the health and sustainability of coral reef ecosystems. Stresses associated with climate change, such as high-temperature episodes that promote coral bleaching, reduced calcification, and changes in ocean and atmospheric circulation, present one set of challenges to coral reefs. However, these stresses may exacerbate other stresses not directly related to climate, such as disease, predation, and the cumulative effects of other nonclimate stresses. Thus, it is difficult to separate the effects of global climate and local nonclimate influences when considering reef condition or vulnerability. 2) Coral reef alteration, degradation, and loss will continue for the foreseeable future, especially in those areas already showing evidence of systemic stress. As we enter an unprecedented climatic state, recent geological and biological history gives us little on which to base predictions regarding the future of coral reef ecosystems. Key uncertainties include the extent to which human activities will continue to alter the environment, how climate variability such as the frequency and intensity of El Niño-Southern Oscillation (ENSO) events will change relative to global temperature, and the biological and ecological responses of coral reef communities to unprecedented future conditions. However, there is no realistic doubt that continued climate change will cause further degradation of coral reef communities, which will be even more devastating in combination with the continuing nonclimate stresses that will almost certainly increase in magnitude and frequency. 3) The effects of climate change on global coral reef ecosystems will vary from one region to another. Although climate change has the potential to yield some benefits for certain coral species in specific regions, such as the expansion of their geographic ranges to higher latitudes, most of the effects of climate change are stressful rather than beneficial. Reef systems that are at the intersection of global climatic and local human stresses will be the most vulnerable. Remote, deep, or well-protected reef communities are more likely to provide reserves and refuges for future generations of coral reef organisms, and aesthetic and scientific resources for future generations of humans. 4) While the net effects of climate change on coral reefs will be negative, coral reef organisms and communities are not necessarily doomed to total extinction. The diversity of coral species comprising existing reefs, the acknowledged adaptation potential of reef organisms, spatial and temporal variations in climate change, and the potential for human management and protection of coral reef ecosystems all provide possibilities for survival. Nevertheless, coral reefs of the future will be fewer and probably very different in community composition than those that presently exist, and these changes will cause further ecological and economic losses. 5) Research into adaptation and recovery mechanisms and enhanced monitoring of coral reef environments will permit us to learn from and influence the course of events rather than simply observe the decline. Most local (and some regional) nonclimate stresses have the potential to be mitigated and

managed more readily than global climate change itself. A significant step would be a distributed international network of coral reef refuges and marine protected areas, selected on the basis of biological and environmental diversity, connectivity, potential threats, and enforcement feasibility. Yet, even with such efforts, recent degradation of coral ecosystems combined with future climate change will still pose a significant challenge to the global sustainability of coral reefs. Burke, L, Selig, L and Spalding, M. (2002) Reefs at risk in Southeast Asia. World Resources Institute: Washington DC (USA), 72 pp. BIOLOGICAL ENDOWMENT Southeast Asia contains nearly 100,000 km2 of coral reefs, almost 34 percent of the world total. With over 600 of the almost 800 reef-building coral species, these reefs have the highest levels of marine biodiversity on earth. Southeast Asia is also the global center of biodiversity for coral reef fish, mollusks, and crustaceans. The region contains 51 of the world’s 70 mangrove species and 23 of the 50 seagrass species. ECONOMIC VALUE The economic value associated with coral reefs in Southeast Asia is substantial. The value of the region’s sustainable coral reef fisheries alone is US$2.4 billion per year. In addition, coral reefs are vital to food security, employment, tourism, pharmaceutical research, and shoreline protection. The coral reefs of Indonesia and the Philippines provide annual economic benefits estimated at US$1.6 billion and US$1.1 billion per year, respectively. THREATS TO REEFS The heavy reliance on marine resources across Southeast Asia has resulted in the overexploitation and degradation of many coral reefs, particularly those near major population centers. The main threats include overfishing, destructive fishing practices, and sedimentation and pollution from land-based sources. Human activities now threaten an estimated 88 percent of Southeast Asia’s coral reefs, jeopardizing their biological and economic value to society. For 50 percent of these reefs, the level of threat is “high” or “very high.” Only 12 percent of reefs are at low risk. The Reefs at Risk project estimates that about 64 percent of the region’s reefs are threatened by overfishing, and 56 percent are threatened by destructive fishing techniques. In addition, dredging, landfilling, mining of sand and coral, coastal construction, discharge of sewage and other activities associated with coastal development threaten about 25 percent of the region’s coral reefs. Sediment and pollution from deforestation and agricultural activities threaten an estimated 20 percent of the region’s reefs. Over 90 percent of the coral reefs in Cambodia, Singapore, Taiwan, the Philippines, Vietnam, China, and the Spratly Islands are threatened, and over 85 percent of the reefs of Malaysia and Indonesia are threatened. Indonesia and the Philippines together possess 77 percent of the region’s coral reefs and nearly 80 percent of all threatened reefs in the region. Logging, destructive fishing practices, overfishing, and other activities that are damaging to coral reefs may be lucrative to individuals in the short-term. However, the net economic losses to society from diminished coastal protection, tourism and sustainable fisheries usually outweigh the short-term benefits. Over a 20-year period, current levels of blast fishing, overfishing, and sedimentation could cost Indonesia and the Philippines more than US$2.6 billion and US$2.5 billion, respectively. Global climate change is also a significant threat to coral reefs in Southeast Asia. Elevated sea-surface temperatures have resulted in more severe and more frequent coral

bleaching. The 1997– 98 El Niño Southern Oscillation (ENSO) event triggered the largest worldwide coral bleaching event ever recorded. In Southeast Asia, an estimated 18 percent of the region’s coral reefs were damaged or destroyed. MANAGEMENT Effective management is key to maintaining coastal resources, but, is inadequate across much of the region. Some 646 marine protected areas (MPAs) include an estimated 8 percent of the region’s coral reefs. Of the 332 MPAs whose management effectiveness could be determined, only 14 percent were rated as effectively managed, 48 percent have partially effective management, and 38 percent have inadequate management. THE LACK OF INFORMATION Despite widespread recognition that coral reefs are severely threatened, information about the status and nature of the threats to specific reef areas is limited. This lack of information inhibits effective decisionmaking concerning coastal resources. The Reefs at Risk project was developed to address this deficiency by creating standardized indicators that raise awareness about threats to coral reefs and highlight the linkages between human activity and coral reef condition. Burke, L and Maidens, J. (2004) Reefs at risk in the Caribbean. World Resources Institute: Washington DC (USA), 80 pp. Coral reefs are an integral part of the Caribbean fabric, threading along thousands of kilometers of coastline. Teeming with fish and invertebrate life, these ecosystems provide food for millions of people. Buffering shorelines, they protect the land from the worst ravages of storms. Coral reefs form the foundation of the thriving Caribbea tourism industry, the region’s most important economic sector. The reefs supply much of the sand for the region’s beautiful beaches and lure divers and snorkelers from far and wide to come and explore the reefs’ colorful and mysterious depths. The dazzling array of species living on coral reefs has also attracted the attention of the pharmaceutical industry as a potential source of new drugs and life-saving medical treatments. Unfortunately, these valuable ecosystems are degrading rapidly under the mounting pressure of many human activities. Coastal development, land clearance, and intensive agriculture all contribute damaging sediment and pollution to coastal waters, while overfishing is changing the ecological balance of coral reef environments. In addition, rising sea temperatures have prompted dramatic “coral bleaching” events in recent years, weakening and killing corals in many areas. At the same time, poorly understood coral diseases have spread rapidly across the region, devastating some of the main reef-building corals. Coral reef degradation and mortality will significantly impact the region’s economy through reduced habitat for fish and shellfish, diminished appeal for tourists, and a lessened capacity to protect the shoreline. Understanding the nature and extent of these threats and their likely economic impacts on the future productivity of Caribbean coral reefs as sources of food, recreation, employment, and biopharmaceuticals is of central importance to conservation and planning efforts. Numerous studies are underway to monitor and assess reef conditions at particular locations in the Caribbean, but data gaps persist and, for the majority of reefs, little information is available. Many such efforts fail to combine ecosystem studies with monitoring of socioeconomic and environmental conditions, making it difficult to link changes in coral condition to specific causes. PURPOSE AND GOAL OF REEFS AT RISK IN THE CARIBBEAN

The Reefs at Risk in the Caribbean project was launched to help protect and restore these valuable, threatened ecosystems by providing decision-makers and the public with information and tools to manage coastal habitats more effectively. The project focuses on compiling, integrating, and disseminating critical information on these precious resources for the entire Caribbean region. This information is intended both to raise awareness about the threats to and value of Caribbean reefs and to encourage greater protection and restoration efforts. Conducted by the World Resources Institute in cooperation with over 20 organizations working in the region, the project represents a unique, region-wide look at the threats facing Caribbean coral reefs. The collaborative process of data gathering and analysis has produced the first regionally consistent, detailed mapping of these threats. The project provides decision-makers and the public with important insights on links between human activities that stress and damage reef organisms and where degradation of reefs could be expected to occur, or may have already occurred. The maps created by the Reefs at Risk project will assist regional and national organizations in setting priorities for conservation and natural resource management. The analytical tools and threat indicators will also allow managers to assess, for the first time, the source and scale of threats affecting those many reef areas for which more detailed monitoring information is unavailable. Bythell, JC, Douglas, AE, Sharp, VA, Searle, JB and Brown, BE. (1997) Algal Genotype and Photoacclimatory Responses of the Symbiotic Alga Symbiodinium in Natural Populations of the Sea Anemone Anemonia viridis. Proceedings: Biological Sciences 264(1386), 1277-1282. As an approach to investigate the impact of solar radiation on an alga-invertebrate symbiosis, tlle genetic variation and photosynthetic responses of the dinoflagellate algal symbiosis in an intertidal and a subtidal populationo f the sea anemone Anemonia viridis were explored. Allozyme analysis of the anemones indicated that the two populations were genetically very similar, with a Nei's index of genetic identity (I) of 0.998. The algae in all animals examined were identified as Symhiodinium of clade a by PCR-RFLP analysis of the small subunit ribosomal RNA gene. 'The symbiosis in the two populations did not differ significantly in algal population density, chlorophyll a content per algal cell or any photosynthetic parameter obtained from studies of the relationship between photosynthesis and irradiance. We conclude that there is not necessarily genetic variation or photosynthetic plasticity of the symbiotic algae in Anemonia viridis inhabiting environments characterized by the different solar irradiances of the subtidal and intertidal habitats. Carpenter, S, Walker, B, Anderies, JM and Abel, N. (2001) From metaphor to measurement: Resilience of what to what? Ecosystems 4, 765-781. Resilience is the magnitude of disturbance that can be tolerated before a socioecological system (SES) moves to a different region of state space controlled by a different set of processes. Resilience has multiple levels of meaning: as a metaphor related to sustainability, as a property of dynamic models, and as a measurable quantity that can be assessed in field studies of SES. The operational indicators of resilience have, however, received little attention in the literature. To assess a system’s resilience, one must specify which system configuration and which disturbances are of interest. This paper compares resilience properties in two

contrasting SES, lake districts and rangelands, with respect to the following three general features: (a) The ability of an SES to stay in the domain of attraction is related to slowly changing variables, or slowly changing disturbance regimes, which control the boundaries of the domain of attraction or the frequency of events that could push the system across the boundaries. Examples are soil phosphorus content in lake districts woody vegetation cover in rangelands, and property rights systems that affect land use in both lake districts and rangelands. (b) The ability of an SES to self-organize is related to the extent to which reorganization is endogenous rather than forced by external drivers. Self-organization is enhanced by coevolved ecosystem components and the presence of social networks that facilitate innovative problem solving. (c) The adaptive capacity of an SES is related to the existence of mechanisms for the evolution of novelty or learning. Examples include biodiversity at multiple scales and the existence of institutions that facilitate experimentation, discovery, and innovation. Carpenter, RC and Edmunds, PJ. (2006) Local and regional scale recovery of Diadema promotes recruitment of scleractinian corals. Ecology Letters 9, 271–280. The phase change from coral to macroalgal dominance on many Caribbean reefs was exacerbated by the mortality of the echinoid Diadema antillarum in 1983–1984, and until recently, this sea urchin has remained rare on reefs throughout the western Atlantic. By the late 1990s, Diadema started to reappear in large numbers on some Jamaican reefs, and by 2000, the high densities were correlated with significantly greater abundances of juvenile corals. Here, we show that dense populations of Diadema now occur over a multi-kilometre-wide scale at six locations scattered along a 4100 km arc across the entire Caribbean. In all cases, these dense populations are found in shallow water (< 6 m depth) on outer reef communities and are associated with reduced macroalgal cover and enhanced coral recruitment. We conclude that population recovery of Diadema is occurring at both local and regional scales, and that grazing by this echinoid is creating conditions favouring the recruitment of corals. Castillo, KD and Helmuth, BST. (2005) Influence of thermal history on the response of Montastraea annularis to short-term temperature exposure. Marine Biology 148, 261-270. Acclimation of reef corals to environmental conditions has been related to metabolic response at large geographic scales, but regional relationships have rarely been described. Physiological responses to temperature increases of Montastraea annularis (Ellis and Solander 1786) from an inner lagoon and an outer barrier reef in the Gulf of Honduras, southern Belize, were compared in May 2003. The hypothesis that inferred differences in thermal history would result in contrasting responses to elevated temperature was tested. Ambient seawater temperatures adjacent to corals at 4–5 m depth were measured every 15 min at inner lagoon and outer barrier reef collection sites for 1 year (June 2002–May 2003). Monthly averages and 3-day running averages (warmest period, July–October 2002) of daily maximum seawater temperatures were significantly higher (by 30.5°C) at inner lagoon reef compared to outer barrier reef sites. M. annularis photosynthesis (P) and respiration (R) rates were measured in respirometers at six temperatures between 29°C and 35°C approximately every hour, with repeated measurements over 3 h. P and R were

significantly lower across most temperature treatments for samples collected from the inner lagoon compared to outer barrier reef. Both inner and outer reef M. annularis displayed an increase in P and R with increasing temperature between 29°C and 32°C, but above 32°C P and R sharply declined. P/R ratio versus temperature showed a significant difference between the elevations of the regression lines suggesting that M. annularis from the outer barrier reefs may have been more physiologically stressed than those from the inner lagoon reefs when exposed to acute temperature changes. These results emphasize that thermal stress must be considered within the context of acclimation temperature, and that short-term exposures may have physiologically important effects on this species. Ceccarelli, DM, Jones, GP and McCook, LJ. (2005) Effects of territorial damselfish on an algal-dominated coastal coral reef. Coral Reefs 24, 606–620. Territorial damselfish are important herbivores on coral reefs because they can occupy a large proportion of the substratum and modify the benthic community to promote the cover of food algae. However, on coastal coral reefs damselfish occupy habitats that are often dominated by unpalatable macroalgae. The aim of this study was to examine whether damselfish can maintain distinctive algal assemblages on a coastal reef that is seasonally dominated by Sargassum (Magnetic Island, Great Barrier Reef). Here, three abundant species (Pomacentrus tripunctatus, P. wardi and Stegastes apicalis) occupied up to 60% of the reef substrata. All three species promoted the abundance of food algae in their territories. The magnitudes of the effects varied among reef zones, but patterns were relatively stable over time. Damselfish appear to readily co-exist with large unpalatable macro algae as they can use it as a substratum for promoting the growth of palatable epiphytes. Damselfish territories represent patches of increased epiphyte load on macroalgae, decreased sediment cover, and enhanced cover of palatable algal turf. Celliers, L and Schleyer, MH. (2002) Coral bleaching on high-latitude marginal reefs at Sodwana Bay, South Africa. Marine Pollution Bulletin 44, 1380-1387. Coral bleaching, involving the expulsion of symbiotic zooxanthellae from the host cells, poses a major threat to coral reefs throughout their distributional range. The role of temperature in coral bleaching has been extensively investigated and is widely accepted. A bleaching event was observed on the marginal high-latitude reefs of South Africa located at Sodwana Bay during the summer months of 2000. This was associated with increased sea temperatures with high seasonal peaks in summer and increased radiation in exceptionally clear water. The bleaching was limited to Two-mile Reef and Nine-mile Reef at Sodwana Bay and affected <12% of the total living cover on Two-mile Reef. Montipora spp., Alveopora spongiosa and Acropora spp. were bleached, as well as some Alcyoniidae (Sinularia dura, Lobophytum depressum, L. patulum). A cyclical increase in sea temperature (with a period of 5–6 years) was recorded during 1998–2000 in addition to the regional temperature increase caused by the El Ni~no Southern Oscillation phenomenon. The mean sea temperature increased at a rate of 0.27°C year-1 from May 1994 to April 2000. High maximum temperatures were measured (>29°C). The lowest mean monthly and the mean maximum monthly temperatures at which coral bleaching occurred were 27.5 and 28.8°C, respectively, while the duration for which high temperatures occurred in 2000 was 67 days at ≥27.5°C (4 days at ≥28.8°C). Increased water clarity and

radiation appeared to be a synergistic cause in the coral bleaching encountered at Sodwana Bay. Cesar, H, Burke, L and Pet-Soede, L. (2003) The economics of worldwide coral reef degradation. Cesar Environmental Economics Consulting: Arnhem (Netherlands), 23 pp. Coral reefs are an incredibly valuable ecosystem. Not only are they very important for nature, but they represent a very high value for humankind, supporting millions of people whose lives depend on these natural resources for a source of food and income. Estimates in this report show that coral reefs provide each year nearly US$ 30 billion in net benefits in goods and services to world economies, including, tourism, fisheries and coastal protection (Table 1). Yet coral reefs are under heavy pressure. Already, 27% is permanently lost and with current trends, a further 30% is at risk of being lost in the coming thirty years. With such devastating levels of destruction, the social and economic implications for the millions of people who depend on coral reefs are of great concern. Over 39% of the world population now live within 100 kilometres of the coast and many people in these areas depend on reefs. Reefs protect coastlines and reef fish provide a source of nutrition and income. Poverty increases and food security decreases as fish stocks are depleted. This drives fishers further toward the use of destructive methods to catch what little there is left. Key causes of coral reef decline have been the over-development of the coastal area and the over-use of coral reef resources. Migration to coastal areas has created a surge in land development leading to clearance of important coastal ecosystems such as mangroves and seagrass beds. Unregulated coastal construction, such as hotels, factories and desalination plants, has increased sedimentation in the coastal waters and is destroying reefs worldwide as light levels in the water column are reduced and reefs are smothered. Untreated sewage and chemical agriculture run-off (e.g. pesticides, herbicides and fertilizers) have caused nutrient loading into coral reef waters, leading to algal blooms and eutrophication. Overfishing and destructive fishing practices have decimated coral reef fish populations and their habitats. In addition, increases in sea surface temperatures associated with global climate change are causing ever more frequent bleaching events. Countries with coral reefs attract millions of SCUBA divers every year, yielding significant economic benefits to the host country. Globally, tourism is estimated to provide US$ 9.6 billion in annual net benefits and a multiple of this amount in tourism spending (Table 1). These revenue streams in coral reef areas are being threatened by the deterioration of coral reefs. Damage to the reefs has often been caused by the increase in tourism itself, through direct damage by careless tourists and through the unregulated construction and the irresponsible operation of tourist related facilities. Mass tourism poses a threat to reefs and to the income that coral reefs provide to the local population. Sustainable tourism, on the other hand, is a source of income to people in reef areas. It even forms an alternative to destructive fishing practices. Reef fisheries provide nutrition and necessary incomes to millions of people in developing countries. Potential reef fishing benefits are estimated at US$ 5.7 billion annually. However, increases in fishing effort, driven by rising populations and more effective fishing technology, have led to overfishing, such that the obtained economic value of coral reef fisheries is now close to zero in many developing countries. With fewer fish on the reefs many fishermen have resorted to more efficient yet highly

damaging methods, such as bomb and cyanide fishing. Because enforcement of these illegal operations is difficult, the financial incentive to fishers in the short run is high. However, the ecological impact causes a net loss to society. For instance, in areas of Indonesia with high potential value of tourism and coastal protection, the socio-economic costs have been estimated to be four times higher than the total net private benefits from blast fishing. The cost of ‘inaction’ on blast fishing has been estimated at US$ 3.8 billion in Indonesia over the last 25 years. These figures would have justified enforcement expenditures of around US$ 400 million annually. Land-based pollution is another major threat to coral reefs as untreated sewage from urban areas and runoff from chemicals used in agriculture cause sedimentation and mass algal growth. Currently 22% of the world’s coral reefs are under medium to high risk from these land-based sources of pollution. As a result, investment to decrease the flow of sediments and excess nutrients into coastal waters is often justified despite high upfront costs. For example, a proposed wastewater treatment plant in the Florida Keys requires around US$ 60-70 million in investment costs and around US$ 4 million annually in operation and mainetance costs. However, in the long-term, the benefits to the local population are much higher, estimated at around US$ 700 million in Net Present Value (NPV) terms. Additional side benefits such as reduction in water-borne diseases provide further arguments in favour of these investments. To be cost-effective, river basin management should be linked with coastal zone management. This is the essence of the H2O Partnership (from Hilltop to Ocean) launched at the World Summit on Sustainable Development in Johannesburg, August 2002. Global climate change and related coral bleaching pose a final set of threats to coral reefs. In 1998, 75% of reefs worldwide were affected by bleaching and 16% suffered mortality. Recent studies predict that bleaching could become an annual event within the next 25 to 50 years. The study here calculates the net present value of future losses over the next 50 years at a 3% discount rate. The losses in the case of the ‘severe’ coral bleaching scenario with mass coral mortality are estimated around US$ 83 billion. For the ‘moderate’ scenario, where bleaching leads to less mortality, costs are estimated at around US$ 21 billion. Other associated impacts of global climate change also have adverse effects on coral reefs, such as increased frequencies of storms and hurricanes. A recent study estimates that climate change will cause losses of US$ 109.9 million in the Caribbean due to increased sea-surface temperatures, sea-level rise and loss of species, among others. This is equal to 13.8% of the total GDP in the region. Reduction of other stressors to reefs such as those mentioned above, in combination with climate change adaptation measures, could help to decrease the impacts of climate change and coral bleaching on reefs. Without even attempting to measure their intrinsic value, it is clear that coral reefs contribute enormously to food, income and various other quantifiable benefits, if properly managed. Net potential benefits are estimated at US$ 30 billion per year. With current trends of reef degradation, revenues will decrease, in some cases by as much as 75% with increasing costs. This will imperil the lives of millions of people who rely on reefs for food and income, especially in developing countries. Good reef management is costly, but the losses will be much higher if we fail to take action now. The choice is ours.

Chabanet, P. (2002) Coral reef fish communities of Mayotte (western Indian Ocean) two years after the impact of the 1998 bleaching event. Marine and Freshwater Research 53, 107-113. This study, conducted during the Coral Reef Observatory programme in 2000, was carried out at 9 localities: 4 on fringing reefs (0 and 3 m depth), 2 on inner reefs (0 and –3 m) and 3 on outer barrier reefs (3 and 6 m). Quantitative data were collected by visual census techniques along 50 m × 5 m transects. The ichthyofauna is separated into fringing inner-reef communities and outer barrier-reef communities. Species richness and abundance were always higher at the deeper sites. These indices were also lower on fringing reef flats and higher on the outer barrier reefs. In terms of individual numbers, browsers of sessile invertebrates (mainly Chaetodontidae) are poorly represented in the fish community at all sites (3%). Planktivores are the dominant trophic category on the deeper sites (mean 63%) and herbivores on the shallower sites (mean 40%, mainly Acanthuridae). The high abundance of herbivores and the low abundance of browsers of sessile invertebrates could be due to altered benthic communities resulting from the 1998 bleaching event. Chen, CA, Wang, J-H, Fang, L-S and Yang, Y-W. (2005) Fluctuating algal symbiont communities in Acropora palifera (Scleractinia: Acroporidae) from Taiwan. Marine Ecology Progress Series 295, 113-121. Seasonal dynamics of algal symbiont communities from the reef flat-dwelling coral Acropora palifera was monitored from January 2000 to July 2001 at Tantzei Bay, Kenting National Park, southern Taiwan. Zooxanthellae density and pigment contents monitored throughout the 18 mo sampling period indicated that no significant bleaching occurred in A. palifera. Molecular phylotyping showed that 2 distinct symbionts, Phylotypes C and D, were associated with A. palifera, either individually or simultaneously. Population surveys throughout the bimonthly sampling period demonstrated that the symbiont community within A. palifera was highly dynamic with significant fluctuations; a drop in the Phylotype D percentage occurred in May, which was correlated with a rise in seawater temperature in the late spring/early summer at the sampled reef. Direct tracking of tagged corals provided evidence that seasonal fluctuations in the algal symbiont communities not only involve changes in zooxanthellae densities and pigment contents, but also a reshuffling of different Symbiodinium phylotypes. Our study highlights that stress tolerance among phylotypes urgently needs to be re-evaluated through a better understanding of the physiological traits of symbionts. Chen, CA, Yang, Y-W, Wei, NV, Tsai, W-S and Fang, L-S. (2005) Symbiont diversity in sceractinian corals from tropical reefs and sub-tropical non-reef communities in Taiwan. Coral Reefs 24, 11-22. We examined zooxanthellae diversity in scleractinian corals from southern Taiwan and the Penghu Archipelago, a tropical coral reef and a subtropical nonreefal community, respectively. Zooxanthellae diversity was investigated in 52 species of scleractinian corals from 26 genera and 13 families, using restriction fragment length polymorphism (RFLP), and phylogenetic analyses of the nuclear small-subunit ribosomal DNA (nssrDNA) and large-subunit ribosomal DNA (nlsrDNA). RFLP and phylogenetic analyses of nuclearencoded ribosomal RNA genes showed that

Symbiodinium clade C was the dominant zooxanthellae in scleractinian corals in the seas around Taiwan; Symbiodinium clade D was also found in some species. Both Symbiodinium clade C and D were found in colonies of seven species of scleractinian corals. Symbiodinium clade D was associated with corals that inhabit either shallow water or the reef edge in deep water, supporting the hypothesis that Symbiodinium clade D is a relatively stress-tolerant zooxanthellae found in marginal habitats. Christie, P, White, A and Deguit, E. (2002) Starting point or solution? Community-based marine protected areas in the Philippines. Journal of Environmental Management 66, 441-454. In 1985, in response to declining coral reef conditions, local residents and officials established small, no-take marine sanctuaries on Balicasag and Pamilacan Islands through a community-based process. The implementation of marine protected areas (MPAs) on Balicasag and Pamilacan Islands has been a partial success. As a direct result of protection, living hard coral cover has increased by 119% in Balicasag's sanctuary and by 67% in the non-sanctuary during the period 1984 to 1999, but Balicasag's reef is increasingly affected by breakage from anchors from dive boats and Crown-of-thorns starfish infestations. During the same period, living hard coral cover decreased by 20% in Pamilacan's sanctuary and by 45% in the non-sanctuary from 1984 to 1999. The decrease in living hard coral cover in Pamilacan's sanctuary is most likely a result of the 1998 bleaching event, Crown-of-thorn starfish and possible storm damage. Although there was an initial increase in the economically important target fish abundance in the Balicasag sanctuary and non-sanctuary and in the Pamilacan sanctuary during the first two years of implementation in the mid-1980s, there has since been a significant decline. Mean target fish abundance for the Balicasag non-sanctuary at 230 (+65) individuals per 500m2 is not significantly different from control sites without MPAs on nearby Panglao and Cabilao Islands at 164 (+67) individuals per 500m2. In general, fish abundance and diversity inside and outside the sanctuaries peaked in 1986, a year after the establishment of the sanctuaries when enforcement was strictest. Therefore, despite considerable success in enforcing regulations associated with these small MPAs at Balicasag and Pamilacan Islands, a trend of declining fish abundance and species richness among economically valuable species immediately outside the no-take areas highlights the limitations of small and isolated MPAs. This study contributes to the growing sentiment that it is not realistic for scattered, small no-take areas to maintain fish abundance and diversity on surrounding reefs when intensive fishing effort immediately adjacent to no-take areas removes most fish that exit these areas. This finding emphasizes the importance of nesting individual MPAs within broader management regimes that lead to overall fishing effort reduction and networking of MPAs. Among other recommendations, the authors advocate for continued support for community-based MPAs, a network of MPAs, reduced fishing effort in areas surrounding the MPAs and other management measures to improve the quality of the coral reef habitats.

Church, J and Obura, DO. (2004) Sustaining coral reef ecosystems and their fisheries in the Kiunga National Reserve, Lamu, Kenya. 10th International Coral Reef Symposium: Okinawa (Japan), 22 pp. Coral reefs in the Kiunga Marine National Reserve (KMNR) (40° 07’ E, 2° 00’ S) are located in a transition ecotone between the warmer East African coral reef bioregion to the south, and colder waters of the Somali Current to the north. The reefs have been monitored annually from 1998 to the present, documenting a range of ecosystem changes from large and small scale threats. Reefs in the area suffered >80% loss of coral cover due to mass bleaching in the 1998 El Niño event, and 25-40% loss of coral species at individual site levels. Recovery of coral community structure has been variable, with some reefs showing strong recovery, while others have declined further. A Harmful Algal Bloom (HAB) and coral disease in early 2002 further impacted these reefs, causing mass mortalities of fish and coral, and failure of coral recruitment in that year. Fishing impacts to the reserve are high, with a strong south-north decline in fish density due to easier access to the migrant and large fishing communities to the south of the reserve. Responsibility for management of the KMNR falls under multiple institutions, including the Kenya Wildlife Service (KWS), Fisheries and Forestry Departments, and the local council. Overlapping mandates, unclear relationships, limited information and understanding, and lack of resources have hampered effective management. The monitoring programme reported here is one aspect of new collaborative approaches to coral reef and fisheries management, and has focused on improving the information and understanding of the biological and resource systems of the area. The ecosystem trends induced by larger scale threats and the south-north fish resource gradient caused by local use patterns will be analyzed in an attempt to develop sustainable management practices for the reserve. Clark, S and Edwards, AJ. (1999) An evaluation of artificial reef structures as tools for marine habitat rehabilitation in the Maldives. Aquatic Conservation: Marine and Freshwater Ecosystems 9, 5-21. 1. In the Maldives, coral mining for the construction industry has resulted in widespread degradation of shallow reef-flat areas. Due to the loss of these coastal resources and the associated problems of coastal erosion, there is an urgent need to find practical methods for rehabilitating mined reefs. 2. The slow rates of natural recovery of mined reefs has prompted interest in the potential of artificial reef structures to rehabilitate these degraded habitats. An experimental artificial reef programme was initiated in 1990 to discover whether it is feasible to use a bio-engineering approach to kick-start natural reef recovery. 3. The main goals of the project were to restore the capacity of degraded reefs for sea defence and their ability to harbour fish species. Accordingly, 360 t of concrete structures of varying levels of topographic complexity, stabilising effect and cost were deployed on a heavily mined study site close to the capital island, Malé. 4. Within 1 year of deployment, the artificial reef structures had similar or greater species richness and densities of reef fish than did control pristine reef flats. However, the community structure of the fish populations on the artificial reef structures was significantly different to that on unmined reef flats. 5. Preliminary results of a monitoring programme indicated that substantial coral recruitment had occurred on the larger reef structures which were each supporting

ca. 500 colonies, some of which were approaching 25 cm in diameter after 3.5 years. An evaluation of the effectiveness of the various artificial reef structures is discussed in relation to their design features and costs and in line with timescales for the recovery processes. Coles, SL and Brown, BE. (2003) Coral bleaching – capacity for acclimatization and adaptation. Advances in Marine Biology 46, 183-223. Coral bleaching, i.e., loss of most of the symbiotic zooxanthellae normally found within coral tissue, has occurred with increasing frequency on coral reefs throughout the world in the last 20 years, mostly during periods of El Niño Southern Oscillation (ENSO). Experiments and observations indicate that coral bleaching results primarily from elevated seawater temperatures under high light conditions, which increases rates of biochemical reactions associated with zooxanthellar photosynthesis, producing toxic forms of oxygen that interfere with cellular processes. Published projections of a baseline of increasing ocean temperature resulting from global warming have suggested that annual temperature maxima within 30 years may be at levels that will cause frequent coral bleaching and widespread mortality leading to decline of corals as dominant organisms on reefs. However, these projections have not considered the high variability in bleaching response that occurs among corals both within and among species. There is information that corals and their symbionts may be capable of acclimatization and selective adaptation to elevated temperatures that have already resulted in bleaching resistant coral populations, both locally and regionally, in various areas of the world. There are possible mechanisms that might provide resistance and protection to increased temperature and light. These include inducible heat shock proteins that act in refolding denatured cellular and structural proteins, production of oxidative enzymes that inactivate harmful oxygen radicals, fluorescent coral pigments that both reflect and dissipate light energy, and phenotypic adaptations of zooxanthellae and adaptive shifts in their populations at higher temperatures. Such mechanisms, when considered in conjunction with experimental and observational evidence for coral recovery in areas that have undergone coral bleaching, suggest an as yet undefined capacity in corals and zooxanthellae to adapt to conditions that have induced coral bleaching. Clearly, there are limits to acclimatory processes that can counter coral bleaching resulting from elevated sea temperatures, but scientific models will not accurately predict the fate of reef corals until we have a better understanding of coral-algal acclimatization/adaptation potential. Research is particularly needed with respect to the molecular and physiological mechanisms that promote thermal tolerance in corals and zooxanthellae and identification of genetic characteristics responsible for the variety of responses that occur in a coral bleaching event. Only then will we have some idea of the nature of likely responses, the timescales involved and the role of 'experience' in modifying bleaching impact.

Conley, J, Harding, S, Barnard, N, Hine, A and Raines, P. (2003) Fiji coral reef conservation project: 1st annual report. Coral Cay Conservation: London (UK), 133 pp. · Much of Fiji’s wealth is generated by its extensive marine resources, which provide, for example, protein based food from fishing and income from tourism. However, a suite of factors currently threatens the ecological balance and health of Fiji’s reef ecosystems. · Following on from the successful pilot project (MCRCP), stakeholders in the Mamanuca Islands invited Coral Cay Conservation (CCC) to continue working in the region, leading to the implementation of a full CCC project entitled ‘Fiji Coral Reef Conservation Project (FCRCP). This report outlines the progress of the FCRCP over the first year of operations (March 2002 – April 2003). · Fieldwork during year one of the FCRCP focused on gathering data from a number of allocated survey sectors over a wide range of geographical locations and habitat types using: baseline transects for habitat mapping and Reef Check surveys to assess reef health. · Results from year one of the FCRCP showed a range of detrimental anthropogenic influences to be present in the Mamanuca Islands. In particular coral bleaching, sedimentation and the combination of nutrient elevation and over exploitation of marine resources are subjects that require attention in the region. Perhaps the most obvious of these impacts was the mass coral bleaching event, which occurred in early 2000, with a second smaller event in 2001. However Reef Check results of hard coral cover indicate that some recovery has taken place (see separate CCC report; Walker et al., 2002). · Of the 31 designated survey sectors, 14 have been completed during the first year of the FCRCP. Preliminary maps showing the habitat composition of survey transects on all surveyed reefs were produced. Analysis of survey data in the form of six analysis areas has revealed a range of habitats in the completed sectors. Habitats with the highest hard coral cover and diversity and most diverse reef fish assemblages have been highlighted. · An extensive environmental awareness and education programme for local communities and stakeholders has successfully been established in the Mamanucas during first year of the FCRCP. It is vital the environmental education goes hand in hand with the CCC marine survey programme so that awareness of the importance of both sides of the FCRCP is known in the region. · The support by many stakeholders for mitigating measures in the Mamanuca Islands represents a clear desire to address the threats to reef health and work towards sustainable use. Such a goal could be addressed by both reducing the threats to reef health and establishing a series of marine reserves. · Marine reserves are important since they: conserve biodiversity; increase fish abundances within the reserve and provide ‘spill-over’ into surrounding areas; facilitate reef recovery; separate conflicting uses; serve as a centre for public education and attract sustainable tourist revenue. · Research indicates that 20% of the reefs of an area should be ‘no-take’ in order to maximise the chances of sustaining the fisheries and given that the reefs delineated on the habitat map cover approximately 70 km2, the eventual aim should be to protect 14 km2 of shallow (<30 m) benthic habitat within the Mamanuca Islands from fishing.

· A number of coral reefs are recommended for the designation of MPAs in the areas surveyed during year one of the FCRCP. These have been selected predominantly on the basis of the analysis of survey data to identify habitats with high hard coral abundance combined with high benthic faunal and reef fish diversity. · The areas recommended for protection are south-east Mana Island, Yalodrivi Reef, Mothui Island, Malolo Patch Reef and Navini Island. The areas selected comprise 3.75 km2 of coral reef habitats leaving 10.25 km2 still awaiting allocation in the remaining time of the FCRCP. · Coral Cay Conservation enters into Year 2 of the FCRCP with four main aims: o To continue baseline surveys in areas that have not been surveyed to allow the production of a full inventory of the marine resources. o To continue and expand community education efforts with all stakeholder groups. o To continue monitoring to examine the recovery of the marine resources of the Mamanucas following episodes of natural and anthropogenic disturbance. o To assist and support any other body or organisation that displays a willingness and interest in the management of the resources of the Mamanuca Islands. Connell, JH, Hughes, TP, Wallace, CC, Tanner, JE, Harms, KE and Kerr, AM. (2004) A long-term study of competition and diversity of corals. Ecological Monographs 74, 179-210. Variations in interspecific competition, abundance, and alpha and beta diversities of corals were studied from 1962 to 2000 at different localities on the reef at Heron Island, Great Barrier Reef, Australia. Reductions in abundance and diversity were caused by direct damage by storms and elimination in competition. Recovery after such reductions was influenced by differences in the size of the species pools of recruits, and in contrasting competitive processes in different environments. In some places, the species pool of coral larval recruits is very low, so species richness (S) and diversity (D) never rise very high. At other sites, this species pool of recruits is larger, and S and D soon rise to high levels. After five different hurricanes destroyed corals at some sites during the 38-year period, recovery times of S and D ranged from 3 to 25 years. One reason for the variety of recovery times is that the physical environment was sometimes so drastically changed during the hurricane that a long period was required to return it to a habitat suitable for corals. Once S and D have peaked during recolonization, they may either remain at a high level, or decline. In shallow water, with no deleterious changes in environmental conditions, S and D may not decline over time, because superior competitors cannot overtop inferior competitors without exposing themselves to deleterious aerial exposure at low tide. At other times and places, S and D did decline over time. One cause of this was a gradual deterioration of the physical environment, as corals grew upward into the intertidal region and died of exposure. S and D also fell because the wave action in hurricanes either killed colonies in whole or part, or changed the drainage patterns over the reef crest, leaving corals high and dry at low tide. At deeper sites, declines in S and D were sometimes caused by heavy wave action, or by interspecific competition, as some corals overgrew or overtopped their neighbors and eliminated them.

Cordio (1999) Coral reef degradation in the Indian Ocean: Status reports and project presentations 1999. Cordio: Stockholm (Sweden), 103 pp. The reports in this volume summarize the extent of damage to the coral reefs of the Indian Ocean caused by the elevated temperatures in 1998. Most of the reports also contain information on the status of reefs 6 to 12 months after the events of 1998, thus describing the fate of dead corals, the first signs of recovery in some areas, and the secondary damage to other organisms dependent on the reef, such as fish. A brief summary of the results is given here. EAST AFRICA • Several areas suffered very high coral mortality. For example, surveys in Kiunga and Malindi (Kenya), Misali and Mafia (Tanzania), and Pemba and Inhacca (Mozambique) showed that 90–100% of the corals died after exposure to water temperatures that exceeded 32°C, mainly in March and April 1998. The mortality of corals culminated around mid-May. In some areas, however, corals continued to die until October. • Following the event in 1998, coral cover appears to have been reduced to between 10 to 50% of previous levels in most areas along the coast of Kenya, Tanzania and northern Mozambique. In some areas the reduction is greater, i.e. up to or above 90%. • Initial investigations indicate that fish communities associated with coral reefs were affected by the coral mortality and that, in general, herbivorous species increased while corallivorous species decreased. • In most affected areas, the cover of algal turf on bleached and dead reefs increased significantly. On Kenyan reefs, for example, algal cover in many areas increased up to 200% as a result of the newly available substrate. INDIAN OCEAN ISLANDS • Coral mortality ranged from 50–90% over extensive areas of shallow reefs in Seychelles, Comoros, Madagascar and Chagos. In some areas (around Mahe, Seychelles), mortality was close to 100%. • By the end of 1998, algal turf covered coral reefs throughout much of the region. • Monitoring of potentially toxic, epiphytic dinoflagellates has shown drastically increased concentrations in areas with dead corals. • By early 1999, much of the dead coral in Chagos was reported to be eroded to rubble, preventing recolonisation. In Socotra Archipelago, coral rubble has been washed ashore and can be found in piles on the beach. • Preliminary assessments of the reef fish communities in Chagos indicate that abundance and diversity have decreased to less than 25% of their former levels. SOUTH ASIA REGION • Bleaching was reported down to a depth of 40 m in Sri Lanka and 30 m in Maldives, as a result of water temperatures of approximately 35°C during the period April to June, 1998. • In Sri Lanka and Maldives, nearly 90% of the corals died in many areas. In the Hikkaduwa and Bar reefs of Sri Lanka, close to 100% of the corals died, and by the end of 1998, these reefs were covered by thick algal turf. In May 1999, large areas of reefs in Maldives showed few signs of recovery. • In India, surveys indicated mortality between 50 and 90% in the reefs in Gulf of Mannar, Andaman Islands and Lakshadweep. • Assessments of the reef fish communities showed drastic reductions in butterfly fish numbers in Sri Lankan reefs.

Cowen, RK, Lwiza, KMM, Sponaugle, S, Paris, CB and Olson, DB. (2000) Connectivity of marine populations: Open or closed? Science 287, 857-859. Most marine populations are thought to be well connected via long-distance dispersal of larval stages. Eulerian and Lagrangian flow models, coupled with linear mortality estimates, were used to examine this assumption. The findings show that when simple advection models are used, larval exchange rates may be overestimated; such simplistic models fail to account for a decrease of up to nine orders of magnitude in larval concentrations resulting from diffusion and mortality. The alternative process of larval retention near local populations is shown to exist and may be of great importance in the maintenance of marine population structure and management of coastal marine resources. Crabbe, MJC and Smith, DJ. (2003) Computer modelling and estimation of recruitment patterns of non-branching coral colonies at three sites in the Wakatobi Marine Park, S.E. Sulawesi, Indonesia; implications for coral reef conservation. Computational Biology and Chemistry 27, 17-27. We have studied growth and estimated recruitment of massive coral colonies at three sites, Kaledupa, Hoga and Sampela, separated by about 1.5 km in the Wakatobi Marine National Park, S.E. Sulawesi, Indonesia. There was significantly higher species richness (PB/0.05), coral cover (PB/0.05) and rugosity (PB/0.01) at Kaledupa than at Sampela. A model for coral reef growth has been developed based on a rational polynomial function. The model was used to simulate typical coral growth curves, and tested using published data obtained by weighing coral colonies underwater in reefs on the south-west coast of Curaçao [‘Neth. J. Sea Res. 10 (1976) 285’]. The model proved an accurate fit to the data, and parameters were obtained for a number of coral species. Surface area data was obtained on over 1200 massive corals at three different sites in the Wakatobi Marine National Park, S.E. Sulawesi, Indonesia. The year of an individual’s recruitment was calculated from knowledge of the growth rate modified by application of the rational polynomial model. The estimated pattern of recruitment was variable, with little numbers of massive corals settling and growing before 1950 at the heavily used site, Sampela, relative to the reef site with little or no human use, Kaledupa, and the intermediate site, Hoga. There was a significantly greater sedimentation rate at Sampela than at either Kaledupa (PB/0.0001) or Hoga (PB/0.0005). The relative mean abundance of fish families present at the reef crests at the three sites, determined using digital video photography, did not correlate with sedimentation rates, underwater visibility or lack of large nonbranching coral colonies. Radial growth rates of three genera of non-branching corals were significantly lower at Sampela than at Kaledupa or at Hoga, and there was a high correlation (r=0.89) between radial growth rates and underwater visibility. Porites spp. was the most abundant coral over all the sites and at all depths followed by Favites (PB/0.04) and Favia spp. (PB/0.03). Colony ages of Porites corals were significantly lower at the 5 m reef flat on the Sampela reef than at the same depth on both other reefs (PB/0.005). At Sampela, only 2.8% of corals on the 5 m reef crest are of a size to have survived from before 1950. The Scleractinian coral community of Sampela is severely impacted by depositing sediments which can lead to the suffocation of corals, whilst also decreasing light penetration resulting in decreased growth and calcification rates. The net loss of material from Sampela, if

not checked, could result in the loss of this protective barrier which would be to the detriment of the sublittoral sand flats and hence the Sampela village. Craig, P, Birkeland, C and Belliveau, S. (2001) High temperature tolerated by a diverse assemblage of shallow-water corals in American Samoa. Coral Reefs 20, 185-189. Corals in shallow waters are subjected to widely fluctuating temperatures on a daily basis. Using continuous temperature recordings, we examined the temperature regime in one such area, a backreef moat with low tide depths of 1-2 m on Ofu Island in American Samoa. The moat supports a high diversity of 85 coral species [H'(log2) = 3.37] with 25-26% live coral coverage. In one section of the moat, a 4,000-m2 pool inhabited by 52 coral species, the mean summer temperature was 29.3°C, but daily temperatures fluctuated up to 6.3 °C and briefly reached a peak of 34.5°C. The duration of hot water events, e.g., ≥32 °C, averaged 2.4 h per event (maximum 5 h) and occurred on 35 summer days, although daily mean temperatures did not exceed 30.5°C and were generally within 0.5°C of that occurring outside the moat at an exposed coastal area. While there was a previous mortality of many acroporids during a long-term (several month) warming period in 1994, at least nine Acropora species and a diverse range of other taxa withstand the current temperature regime. Craig, P and Basch, L. (2001) Developing a coral reef monitoring program for the national park of American Samoa: A practical, mangement-driven approach for small Marine Protected Areas. Proceedings of Vital Signs Workshop: Pago Pago (American Samoa), 16 pp. The National Park of American Samoa has three small marine protected areas in the Samoan Archipelago, central South Pacific Ocean. At present, the Park is in an early stage of development with low visitation by tourists; Park uses are primarily subsistence fishing by local villagers. A 'Vital Signs' workshop was held to develop a coral reef monitoring program for the Park, where the intent was to focus on the most significant indicators (i.e., the vital signs) of long-term ecological trends or priority concerns of the Park. The workshop first identified who the monitoring program was for (the on-site manager) and then examined what information the manager needed and why. That, in turn, required knowledge of natural environmental changes and threats to the Park's reefs, which are presently local fishing pressure and natural forces (e.g., hurricanes, increased mortalities due to global warming). Monitoring questions related to these natural changes and threats were formulated, followed by a listing of indicators that would provide the desired information by tracking changes in reef condition over time. The vital signs selected were human uses of the park and selected parameters for corals, other invertebrates, fish, algae, and water quality. Vital signs were identified that should be measured regularly (at least yearly), or less frequently to document baseline conditions for possible future comparisons. This vital signs approach provided a convenient way to identify, organize and prioritize the variables that should be included in a monitoring plan. It also identified some needs that exceed current Park capabilities. The next steps in this process will be to determine appropriate sampling designs and methods for the monitoring program.

Crosby, MP, Brighouse, G and Pichon, M. (2002) Priorities and strategies for addressing natural and anthropogenic threats to coral reefs in the Pacific Island Nations. Ocean and Coastal Management 45, 121-137. Marine habitats and resources have been assumed to be almost unlimited, and that if one habitat became degraded or a particular fisheries resource depleted, there always would be another to replace it. The importance of coral reef ecosystems, in particular, may be seen intheir numerous ecological, aesthetic, economic and cultural functions. Pacific islanders recognize that healthy reefs are essential for creation, support, protection and repair of their islands, and serve as a living pantry for the subsistence harvest and consumption of many reef organisms. The ability of coral reef ecosystems to exist in balanced harmony with other naturally occurring competing/limiting physico-chemical and biological agents has been severely challenged in the last several decades by the dramatically increased negative and synergistic impacts from poorly managed anthropogenic activities. In addressing these threats, a paradigm shift may be occurring in the evolution of the role of scientists in society from simply observers of the natural world with tenuous linkages to resource managers and the public, to partners in modern society’s quest for answers to pressing questions related to sustainable use and conservation of coral reef resources. Management principles are beginning to include human motivation and responses as part of coral ecosystems being studied and managed. Managers of coral reef resources face the challenge of balancing conservation and development objectives in the context of the inherent uncertainty of natural systems and the political and social pressures of human systems. Working together, scientists, managers and policymakers can develop priorities and strategies for societal and economic decisions that are strongly coupled with an increasingly comprehensive understanding of the environment. This in turn will lead to both socio-economic health and coral ecosystem health. Employing this new paradigm for interactions between scientists, managers and policymakers, participants of the Pacific Regional ICRI Symposium in Noumea, New Caledonia developed three principal sets of recommendations for addressing natural and anthropogenic threats to coral reefs in Pacific Island Nations: (1) develop and implement a new overarching Participatory Island Ecosystem Management System (PIEMS) for each of the Pacific Island Nations; (2) improve existing, as well as design and implement new, capacity-building programs; and (3) improve scientific understanding of coral reef ecosystems with effective translation and transfer of information. Details for each set of recommendations are provided in this paper. Day, JC. (2002) Zoning – Lessons learnt from the Great Barrier Reef Marine Park. Ocean and Coastal Management 45, 139-156. The Great Barrier Reef Marine Park (GBRMP) is bigger than the United Kingdom, Holland and Switzerland combined. Over the last 25 years a range of management ‘tools’, including zoning plans, permits, education, and more recently management plans, have been applied to regulate access and to control and mitigate impacts associated with human use of the GBRMP. A multiple-use zoning approach provides high levels of protection for specific areas whilst allowing reasonable uses, including certain fishing activities, to continue in other zones. Zoning has long been regarded as a cornerstone of Marine Park management, separating conflicting uses through application of the various zones and determining the appropriateness of various

activities. Zoning in the GBRMP has evolved and changed considerably since the first zoning plan in 1981, along with other management approaches. This paper outlines what aspects of zoning have worked well, what has necessarily changed, and the zoning lessons learned from over two decades of ‘adaptive management’. D’Croz, L and Maté, JL. (2004) Experimental responses to elevated water temperature in genotypes of the reef coral Pocillopora damicornis from upwelling and non-upwelling environments in Panama. Coral Reefs 23, 473-483. The authors investigated the response to experimentally elevated water temperature in genotypes of Pocillopora damicornis from three coral reefs in the upwelling Gulf of Panama and four coral reefs in the non-upwelling Gulf of Chiriquí, Panamanian Pacific. Sea-surface temperature in the Gulf of Panama declines below 20 °C during seasonal upwelling, while in the thermally stable Gulf of Chiriquí, the temperature ranges from 27 to 29 °C. Genotypes of P. damicornis from the seven locations were determined by allozyme electrophoresis. The most abundant genotype at each location was selected for a thermal tolerance experiment where corals were exposed to water temperature of 30 °C (1 °C above ambient) for 43 days. Four site coral genotypes can be uniquely differentiated by the GPI locus, two by the LGG-2 locus, and two by a combination of the MDH-1, LGG-2, and LTY-3 loci. A visual assessment of the coral condition after exposure to an elevated temperature showed that corals from localities in the non-upwelling environment retained a normal to slightly pale appearance, while corals from the upwelling environment bleached and their polyps were mostly retracted. A two-way ANOVA confirmed that corals were significantly affected by water temperature and locality. The zooxanthellae were also significantly affected by the interaction of elevated temperature and locality of the corals. Mean zooxanthellae density decreased by 25 and 55%, respectively, in experimentally heated corals from the non-upwelling and upwelling environments. Low concentrations of photosynthetic pigments per live area of the corals were the norm in corals under elevated temperature. The mean concentration of chlorophyll a per live area of the corals was reduced by 17 and 49%, respectively, in heated corals from the non-upwelling and upwelling sites. Coral genotypes from the upwelling Gulf of Panama demonstrated higher vulnerability to thermal stress than coral genotypes from the non-upwelling Gulf of Chiriquí. However, the latter showed greater differences in their responses. Thus, even at small geographic scales, corals can display different levels of tolerance to thermal stress. The difference in thermal tolerance between corals from upwelling and non-upwelling environments is concomitant with greater genetic differences in experimental corals from the thermally stable Gulf of Chiriquí compared with corals from the upwelling Gulf of Panama. Dizon, RT and Yap, HT. (2005) Effects of coral transplantation in sites of varying distances and environmental conditions. Marine Biology. Several scleractinian coral species with different growth forms and life history strategies were studied in terms of colony growth (expressed as projected linear increment) and survivorship over a range of distances and environmental conditions in the Philippines. The experimental design consisted of 1 m2 plots grouped within a reef site, to several sites within reef systems separated by a distance of about 340

km. There were distinct differences among species, with submassive and massive forms displaying slower growth but better survival, confirming results of other studies. They probably play the role of framework builders of the reef. In contrast, the delicate branching and foliose species had higher growth rates but poor survivorship. This observation, plus their ease of fragmentation, suggests they act more as fillers of the reef matrix. There was high variability in colony increment of a species among the square meter plots, but not among sites within a reef system. Thus, more regular pattern could be observed at this level. In contrast to growth, survivorship differed significantly among sites, being lowest in the site which harbored the greatest amount of dead coral. Growth and survival, however, are not sufficient performance measures to evaluate the success of coral transplantation. Reproduction and subsequent recruitment must also be taken into account. It is recommended that coral transplant and restoration studies consider the broad environmental context of restoration and seek to develop assembly rules that will allow practitioners to match coral types and sequence of interventions to each unique context. Done, T. (2000) Coral reefs and global change: Adaptation, acclimation or extinction? Coral reefs and global change workshop and symposium: Boston (USA). Major revisions of concepts about corals and reef systems were developed by an international working group of scientific experts that met in conjunction with the Society for Integrative and Comparative Biology, the International Society for Reef Studies, and the Ecological Society of America (Boston, January 3-11, 1998) to evaluate the scientific basis for growing concerns about the survival of coral reef ecosystems facing global change and local stresses. The group, sponsored by the Scientific Committee on Oceanic Research (SCOR) and the Land-Ocean Interactions in the Coastal Zone (LOICZ) core project of the International Geosphere-Biosphere Programme (IGBP), and with the support of the NOAA Coastal Ocean Program, produced an interdisciplinary synthesis with important implications for research, assessment, and management. Key conclusions were:

1. The calcification rates of corals, coralline algae, and coral-algal communities depends on the calcium carbonate saturation state of surface seawater, and are expected to be reduced by rising atmospheric carbon dioxide. This represents a global, systemic, climate-related threat to the functioning of reef ecosystems that will interact with the more immediate anthropogenic local stresses.

2. Coral reefs and communities are products of processes operating over a wide range of interacting time and space scales, with fundamentally different controls operating at different scales. While short-term responses will be controlled by local environmental conditions and biotic responses, the longer-term sustainability of a reef system depends on the recruitment, dispersal, persistence, and interactions of populations at larger scales.

3. Corals, and to some extent reef communities, possess numerous mechanisms for acclimatization and adaptation -- diverse reproductive strategies, flexible symbiotic relationships, physiological acclimatization, habitat tolerance, and a range of community interactions. However, current understanding of these mechanisms, as well as of the critically important calcification mechanisms, is inadequate to explain the past success of corals and reefs or to ensure their conservation for the future.

Unlike many terrestrial ecosystems, coral reef ecosystems appear to be directly threatened by globally increasing atmospheric CO2. Therefore, conservation or management strategies aimed at removing or mitigating only local, human-derived, or recently applied environmental stresses are likely to be inadequate. Corals and reefs are potentially robust and resilient, but realizing that potential requires the development of new approaches and greater integration of fundamental and applied research, conservation, and management. Done, T, Turak, E, Wakeford, M, Wooldridge, S, Kininmonth, S, De’ath, G, Berkelmans, R and van Oppen, M. (2004) Testing bleaching resistance hypotheses for the 2002 Great Barrier Reef bleaching event: A contribution to the Transforming Coral Reef Conservation initiative and toolkit. Australian Institute of Marine Science: Townsville (Australia), 11 pp. Selection of areas for protection within coral reef MPAs could benefit from a better understanding of factors that confer resistance to coral bleaching and mortality. Suggested factors include regional oceanographic settings and local topography that promote local cooling of the water, and the species and genotypic composition of the local coral/zooxanthellae community. We assessed the 2002 bleaching event on the Great Barrier Reef and Coal Sea in light of these suggestions. The aims of the study were both to understand how factors interacted to promote areas of good survival in the Great Barrier Reef, and to develop effective scientific process and approach that could be applied elsewhere and promulgated through a toolkit for MPA practitioners, being developed by The Nature Conservancy. We assembled a very strong data-set, including: our ecological and taxonomic assessments over about half the length of the Great Barrier Reef; daily remote sensed sea surface temperatures (SST) for the summer of 2001-2; historical SST data; oceanographic data; bathymetry. All were managed within AIMS Geographic Information System. We used a Bayesian Belief Network (BBN) as a framework to codify some prior beliefs and investigate dependencies among a series of proxies for the potential drivers and responses of bleaching: the remotely sensed environmental stress (sea surface temperature - SST); the geographic setting; and topographic and ecological attributes of reef sites for which we had field data on bleaching impact. Sensitivity analyses helped us to refine and update our beliefs in a manner that improved our capacity to hindcast areas of high and low bleaching impact. Our best predictive capacity came by combining proxies for a site’s heat stress in 2002 (remotely sensed), acclimatization temperatures (remote sensed), the ease with which it could be cooled by tidal mixing (modelled), and type of coral community present at a sample of survey sites (field data). Done, T, Harriott, V, Berkelmans, R and Goggin, L. (2005) Coral bleaching and global climate change: Current state of knowledge. CRC Reef Research Centre: Townsville (Australia). Coral bleaching is the loss of colour from corals under stressful environmental conditions. While several sources of stress can cause corals to bleach, unusually high water temperature and light intensity has been the major cause of coral bleaching events worldwide in recent decades. In 1998 and 2002, major bleaching events in waters of the Great Barrier Reef raised concerns about the health of the

reef. With predictions that temperatures will continue to increase as a result of global climate change, there is growing concern about the future of tropical coral reefs. Done, T, Turak, E, Wakeford, M, DeVantier, L, McDonald, A and Fisk, D. (2007) Decadal changes in turbid-water coral communities at Pandora Reef: loss of resilience or too soon to tell? Coral Reefs. Coral communities were monitored at Pandora Reef, nearshore Great Barrier Reef from 1981 to 2005 using photography and videography. In the 1980s, regional elevation of land-based nutrients in coastal waters (ca. 2–6 times pre-European levels of early 1800s) did not prevent overall recovery of coral cover and diversity following a sequence of environmental disturbances in the 1970s. However, prospects for a repeat of such resilience following catastrophic mortality from high-temperature bleaching in 1998 and a cyclone in 2000 are not clear. Different coral communities around the reef varied greatly in relation to impacts and recovery. Fore-reef communities dominated by acroporids (fast growing branching and tabular Acropora and foliose Montipora) recovered strongly in the 1980s following apparently severe impacts by cyclone, flood and heat wave disturbances in the 1970s, attaining 60–90% cover by stabilizing rubble and outgrowing macro-algae in <10 years. In the back-reef, by contrast, poritid-dominated communities (massive and finger Porites and columnar Goniopora and Alveopora) had more stable trajectories and smaller impact from recent disturbances: recovery was well underway in 2005. The contrasting trajectories of different parts of the reef reflect differential survival of more persistent versus more ephemeral taxa, notably poritids and acroporids, respectively, both major contributors to framework and cover on reefs globally. A repeat of earlier resilience appears possible in the shallow fore-reef, but unlikely in the deeper fore-reef, which had few viable fragments or recruits in 2005. The main limits on recovery may be (1) reduced supply of coral larvae due to widespread regional losses of coral brood stock and (2) the reduced intervals between disturbances associated with global climate change. The presence of a high abundance of Acroporidae is a major pre-disposing risk factor for climate change impacts. Donner, SD, Skirving, WJ, Little, CM, Oppenheimer, M and Hoegh-Guldberg, O. (2005) Global assessment of coral bleaching and required rates of adaptation under climate change. Global Change Biology 11, 2251-2265. Elevated ocean temperatures can cause coral bleaching, the loss of colour from reefbuilding corals because of a breakdown of the symbiosis with the dinoflagellate Symbiodinium. Recent studies have warned that global climate change could increase the frequency of coral bleaching and threaten the long-term viability of coral reefs. These assertions are based on projecting the coarse output from atmosphere–ocean general circulation models (GCMs) to the local conditions around representative coral reefs. Here, we conduct the first comprehensive global assessment of coral bleaching under climate change by adapting the NOAA Coral ReefWatch bleaching prediction method to the output of a low- and high-climate sensitivity GCM. First, we develop and test algorithms for predicting mass coral bleaching with GCM-resolution sea surface temperatures for thousands of coral reefs, using a global coral reef map and 1985–2002 bleaching prediction data. We then use the algorithms to determine the frequency of coral bleaching and required thermal adaptation by corals and their endosymbionts under two different emissions

scenarios. The results indicate that bleaching could become an annual or biannual event for the vast majority of the world’s coral reefs in the next 30–50 years without an increase in thermal tolerance of 0.2–1.0°C per decade. The geographic variability in required thermal adaptation found in each model and emissions scenario suggests that coral reefs in some regions, like Micronesia and western Polynesia, may be particularly vulnerable to climate change. Advances in modelling and monitoring will refine the forecast for individual reefs, but this assessment concludes that the global prognosis is unlikely to change without an accelerated effort to stabilize atmospheric greenhouse gas concentrations. Douglas, AE. (2003) Coral bleaching – How and why? Marine Pollution Bulletin 46, 385-392. Bleaching refers to the loss of colour in symbioses between dinoflagellate algae of the genus Symbiodinium and marine benthic animals, e.g. corals. Bleaching generally results in depressed growth and increased mortality, and it can be considered as a deleterious physiological response or ailment. An explanatory framework for the causes of bleaching comprises three elements: the external factors or triggers of bleaching, e.g. elevated temperature; the symptoms, including elimination of algal cells and loss of algal pigment; and the mechanisms, which define the response of the symbiosis to the triggers, resulting in the observed symptoms. The extent to which bleaching in different symbioses and in response to different triggers involves common mechanisms is currently unknown, but a contribution of interactions between the algal and animal partners to bleaching is predicted. Symbioses vary in their susceptibility to bleaching as a result of genetic variation in Symbiodinium and acclimatory responses of the animal. The evolutionary explanation for bleaching is obscure. Perhaps, bleaching was of selective advantage to the animal hosts under different (more benign?) environmental conditions than the present, or bleaching may be a negative by-product of an otherwise advantageous symbiotic trait, such as the elimination of damaged algal cells. Downing, N, Buckley, R, Stobart, S, LeClair, L and Teleki, K. (2003) Reef fish diversity at Aldabra atoll, Seychelles, during the five years following the 1998 coral bleaching event. Philosophical Transactions: Mathematical, Physical and Engineering Sciences. Quantitative surveys of fish-species diversity were undertaken at 10 m and 20 m water depth on the outer reef at Aldabra Atoll, southern Seychelles, between November 1999 and May 2003. No significant changes in total fish-species diversity, numbers of families represented by these species, or numbers of pomacentrid or chaetodontid species were seen, contrary to fish-diversity changes seen on coral bleaching-impacted reefs elsewhere. The lack of additional anthropogenic pressures at remote Aldabara may make this system, and others like it, more tolerant of bleaching-related population changes.

Downs, CA, Mueller, E, Phillips, S, Fauth, FE and Woodley, CM. (2000) A molecular biomarker system for assessing the health of coral (Montastrea faveolata) during heat stress. Marine Biotechnology 2, 533-544. Using a novel molecular biomaker system (MBS), we assessed the physiological status of coral (Montastraea faveolata) challenged by heat stress by assaying specific cellular and molecular parameters. This technology is particularly relevant for corals because heat stress is thought to be an essential component of coral bleaching. This phenomenon is widely believed to be responsible for coral mortality worldwide, particularly during 1997–1998. Specific parameters of coral cellular physiology were assayed using the MBS that are indicative of a nonstressed or stressed condition. The MBS distinguished the separate and combined effects of heat and light on the 2 coral symbionts, a scleractinian coral and a dinoflagellate algae (zooxanthellae). This technology aids in the accurate diagnosis of coral condition because each parameter is physiologically well understood. Finally, the MBS technology is relatively inexpensive, easy to implement, and precise, and it can be quickly adapted to a high-throughout robotic system for mass sample analysis. Downs, CA, Fauth, JE, Halas, JC, Dustan, P, Bemiss, J and Woodley, CM. (2002) Oxidative stress and seasonal coral bleaching. Free Radical Biology and Medicine 33, 533-543. During the past two decades, coral reefs have experienced extensive degradation worldwide. One etiology for this global degradation is a syndrome known as coral bleaching. Mass coral bleaching events are correlated with increased sea-surface temperatures, however, the cellular mechanism underlying this phenomenon is uncertain. To determine if oxidative stress plays a mechanistic role in the process of sea-surface temperature-related coral bleaching, we examined corals along a depth transect in the Florida Keys over a single season that was characterized by unusually high sea-surface temperatures. We observed strong positive correlations between accumulation of oxidative damage products and bleaching in corals over a year of sampling. High levels of antioxidant enzymes and small heat-shock proteins were negatively correlated with levels of oxidative damage products. Corals that experienced oxidative stress had higher chaperonin levels and protein turnover activity. Our results indicate that coral bleaching is tightly coupled to the antioxidant and cellular stress capacity of the symbiotic coral, supporting the mechanistic model that coral bleaching (zooxanthellae loss) may be a final strategy to defend corals from oxidative stress. Dulvy, NK, Freckleton, RP and Polunin, NVC. (2004) Coral reef cascades and the indirect effects of predator removal by exploitation. Ecology Letters 7, 410-416. Fisheries exploitation provides the opportunity to examine the ecosystem-scale biodiversity consequences of predator removal. We document predatory reef fish densities, coral-eating starfish densities and coral reef structure along a 13-island gradient of subsistence exploitation in Fiji. Along the fishing intensity gradient, predator densities declined by 61% and starfish densities increased by three orders of magnitude. Reefbuilding corals and coralline algae declined by 35% and were replaced by non-reef building taxa (mainly filamentous algae), as a result of starfish

predation. Starfish populations exhibited thresholds and Allee-type dynamics: population growth was negative under light fishing intensities and high predator densities, and positive on islands with higher fishing intensities and low predator densities. These results suggest the depletion of functionally important consumer species by exploitation can indirectly influence coral reef ecosystem structure and function at the scale of islands. Dunlap, WC and Shick, JM. (1998) Ultraviolet radiation-absorbing mycosporine-like amino acids in coral reef organisms: A biochemical and environmental perspective. Journal of Phycology 34, 418-430. Abstract not available. Dunne, RP and Brown, BE. (2001) The influence of solar radiation on bleaching of shallow water reef corals in the Andaman Sea, 1993-1998. Coral Reefs 20, 201-210. Historically, elevated sea-temperature excursions have been used almost exclusively to explain incidences of natural coral bleaching. Little attention, however, has been paid to instances where environmental conditions appear conducive to bleaching but none has been observed. In this paper we examine contemporaneous records of sea temperature and solar radiation (photosynthetically active radiation; PAR, 400-700 nm) over a 6-year period at a site in the Andaman Sea where regular monitoring of shallow water reef coral communities has been conducted since 1979. Four years (1991, 1995, 1997, and 1998) when anomalous sea temperatures were recorded in May are compared. We suggest a complex interaction of PAR and sea temperature, whereby elevated solar radiation prior to sea-temperature maxima may actually protect corals against subsequent bleaching. In addition, we demonstrate the important role that sea-level anomalies play in modifying the underwater light regime to bring about such conditions. Edmunds, PJ and Carpenter, RC. (2001) Recovery of Diadema antillarum reduces macroalgal cover and increases abundance of juvenile corals on a Caribbean reef. Proceedings of the National Academy of Sciences 98, 5067-5071. The transition of many Caribbean reefs from coral to macroalgal dominance has been a prominent issue in coral reef ecology for more than 20 years. Alternative stable state theory predicts that these changes are reversible but, to date, there is little indication of this having occurred. Here we present evidence of the initiation of such a reversal in Jamaica, where shallow reefs at five sites along 8 km of coastline now are characterized by a sea urchin-grazed zone with a mean width of 60 m. In comparison to the seaward algal zone, macroalgae are rare in the urchin zone, where the density of Diadema antillarum is 10 times higher and the density of juvenile corals is up to 11 times higher. These densities are close to those recorded in the late 1970s and early 1980s and are in striking contrast to the decade-long recruitment failure for both Diadema and scleractinians. If these trends continue and expand spatially, reefs throughout the Caribbean may again become dominated by corals and algal turf.

Edmunds, PJ, Gates, RD and Gleason, DF. (2001) The biology of larvae from the reef coral Porites astreoides, and their response to temperature disturbance. Marine Biology 139, 981-989. Pelagic larvae play a fundamental role in the life history of virtually all scleractinian corals, yet much of their biology remains unexplored. One aspect of coral larvae - their response to temperature perturbations -has potentially important consequences for understanding the effects of global warming and El Niño-Southern Oscillation events on coral recruitment. The present study tests the effects of temperature perturbations on coral larvae using Porites astreoides larvae as a model system. In June 1999, larvae were collected from 18 m depth on Conch Reef, Florida, and incubated at ambient (28°C), depressed (26°C), and elevated (33°C) temperatures in outdoor tanks shaded from direct sunlight. Treatments were repeated with new larvae every 24 h, and treatment effects quantified as larval motility, mortality, metamorphosis and metabolism. Elevated temperature significantly increased mortality and metamorphosis, and a similar trend was observed at the reduced temperature, although this was not significant. Neither elevated nor reduced temperatures affected larval motility. Gross photosynthesis (P) was significantly depressed by elevated and reduced temperatures, and respiration (R) varied proportionately with temperature (Q10≈2), although this effect was not statistically significant. At the highest temperature the P/R ratio declined to < 1, indicating that thermal stress reduces the potential for autotrophy. Together, these results suggest that elevated temperatures affect coral larvae by depressing photosynthesis and creating an energy shortage, which ultimately could reduce recruitment (by increasing mortality), shorten larval longevity and favor premature metamorphosis. An unexpected finding was that larvae differed physiologically among release dates. Although preliminary, this suggests that larval fitness in Porites spp. may vary depending on the day of release, a phenomenon that could have significant ramifications with respect to the population structure of adults. Edmunds, PJ. (2002) Long-term dynamics of coral reefs in St. John, US Virgin Islands. Coral Reefs 21, 357-367. In this study, long-term (1987–1998) dynamics are described on a local scale (<20 km) for coral reefs in St. John, US Virgin Islands, which are located in a marine protected area (MPA). The study consists of two sites (Yawzi and Tektite) which were selected in 1987 based on relatively high coral cover (=32%), and six sites that were randomly selected in 1992. Over 12 years, mean coral cover at Yawzi (9 m depth) changed significantly, declining from 45 to 20% cover between 1987 and 1998 (a 56% reduction). Less than 1 km away at Tektite (14 m depth), coral cover also changed significantly, but here it increased 34% (from 32 to 43% cover). Over the same period, macroalgal cover showed a significant upward trend at both sites, increasing from 2 to 26% at Yawzi, and from 6 to 13% at Tektite. The random sites (7–9 m depth) differed from the initial sites in both community structure and dynamics. Mean coral cover at the random sites (~8%) was less than one third of that at Yawzi and Tektite, and varied significantly among sites and years in an idiosyncratic pattern. The percentage cover of macroalgae and the pooled coverage of crustose coralline algae, algal turf, and bare space showed a strong site · time interaction, illustrating that the sites differed in dynamics, but that the differences varied among times. Thus, as has been reported elsewhere in the Caribbean, serious

reef degradation has occurred on at least one reef in St. John, but the patterns of change vary markedly on a kilometer-wide scale. In comparison with other long-term studies of Caribbean coral reefs, the degradation of a coral reef in an MPA around St. John is noteworthy since there are few local anthropogenic disturbances that can be held responsible for the decline. The strong possibility that large-scale events such as hurricanes and global warming have played a pivotal role in the decline of at least one reef in St. John emphasizes the need to embrace landscape- and regional-scale phenomena in order to understand and manage local coral reef dynamics. The occurrence of small patches of relatively healthy reef (i.e., at Tektite) appears trivial in comparison to region-wide reef decline, but such anomalies should be studied further because of their potential roles as refugia for corals and reef-associated taxa. Edmunds, PJ, Gates, RD and Gleason, DF. (2003) The tissue composition of Montastrea franksi during a natural bleaching event in the Florida Keys. Coral Reefs 22, 54-62. In this study, we used a correlative approach to (1) test for an association between bleaching and host tissue composition during a natural bleaching event, and (2) assess whether bleaching susceptibility varies between years. In August 1997, Montastraea franksi at 15-m depth on Conch Reef, Florida, bleached and the severity of the response varied among individuals. Seventy-five randomly selected colonies were quantified for bleaching using both an ordinal scale, assigned by eye, and a continuous scale, assessed using red, green, and blue (RGB) spectral analysis of photographs. Zooxanthella density and chlorophyll a content were evaluated as measures of bleaching, and coral tissue was analyzed for glycerol, free amino acids (FAA), protein, and mycosporine-like amino acids (MAAs); collectively, these are described as the ‘‘tissue composition.’’ In 1998, most of the same coral colonies were analyzed for color and zooxanthella density to determine whether colony color in 1997 was correlated with color in 1998. In 1997, colonies of M. franksi that were ranked by color differed significantly in RGB brightness, zooxanthella density, and chlorophyll a content, but not in tissue composition. Similarly, a multivariate test for a linear relationship between color and tissue composition did not reveal a significant association. Analyses of corals in 1997 and 1998 revealed a significant positive relationship between color in both years (i.e., the same colonies were similarly colored in each year). These results are discussed in the context of the temporal scale of the sampling regime, the nature of the measured traits, and the adaptive bleaching hypothesis. Edmunds, PJ. (2005) The effect of sub-lethal increases in temperature on the growth and population trajectories of three scleractinian corals on the southern Great Barrier Reef. Oecologia 146, 350–364. To date, coral death has been the most conspicuous outcome of warming tropical seas, but as temperatures stabilize at higher values, the consequences for the corals remaining will be mediated by their demographic responses to the sub-lethal effects of temperature. To gain insight into the nature of these responses, here I develop a model to test the effect of increased temperature on populations of three pocilloporid corals at One Tree Island, near the southern extreme of the Great Barrier Reef (GBR). Using Seriatopora hystrix, S. caliendrum and Pocillopora damicornis as study species, the effects of temperature on growth were determined empirically, and the

dynamics of their populations determined under natural temperatures over a 6-month period between 1999 and 2000 [defined as the study year (SY)]. The two data sets were combined in a demographic test of the possibility that the thermal regime projected for the southern GBR in the next 55–83 years—warmer by 3°C than the study year (the SY+3 regime), which is equivalent to 1.4°C warmer than the recent warm year of 1998—would alter coral population trajectories through the effects on coral growth alone; the analyses first were completed by species, then by family after pooling among species. Laboratory experiments showed that growth rates (i.e., calcification) varied significantly among species and temperatures, and displayed curvilinear thermal responses with growth maxima at ~27.1°C. Based on these temperature-growth responses, the SY+3 regime is projected to: (1) increase annualized growth rates of all taxa by 24–39%, and defer the timing of peak growth from the summer to the autumn and spring, (2) alter the intrinsic rate of population growth (k) for S. hystrix (k decreases 26%) and S. caliendrum (k increases 5%), but not for P. damicornis, and (3) have a minor effect on k (a 0.3% increase) for the Pocilloporidae, largely because k varies more among species than it does between temperatures. Ten-year population projections suggest that the effects of a sub-lethal increase in temperature (i.e., the SY+3 regime) are relatively small compared to the interspecific differences in population dynamics, but nevertheless will alter the population size and increase the relative abundance of large colonies at the expense of smaller colonies for all three species, as well as the Pocilloporidae. These effects may play an important role in determining the nuances of coral population structure as seawater warms, and their significance may intensity if the coral species pool is depleted of thermally sensitive species by bleaching. Edwards, AJ and Clark, S. (1998) Coral transplantation: A useful management tool or misguided meddling? Marine Pollution Bulletin 37, 474-487. The primary objectives of coral transplantation are to improve reef `quality' in terms of live coral cover, biodiversity and topographic complexity. Stated reasons for transplanting corals have been to: (1) accelerate reef recovery after ship groundings, (2) replace corals killed by sewage, thermal e.uents or other pollutants, (3) save coral communities or locally rare species threatened by pollution, land reclamation or pier construction, (4) accelerate recovery of reefs after damage by Crown-of-thorns starfish or red tides, (5) aid recovery of reefs following dynamite fishing or coral quarrying, (6) mitigate damage caused by tourists engaged in water-based recreational activities, and (7) enhance the attractiveness of underwater habitat in tourism areas. Whether coral transplantation is likely to be effective from a biological standpoint depends on, among other factors, the water quality, exposure, and degree of substrate consolidation of the receiving area. Whether it is necessary (apart from cases related to reason 3 above), depends primarily on whether the receiving area is failing to recruit naturally. The potential benefits and dis-benefits of coral transplantation are examined in the light of the results of re- search on both coral transplantation and recruitment with particular reference to a 4.5 year study in the Maldives. We suggest that in general, unless receiving areas are failing to recruit juvenile corals, natural recovery processes are likely to be sufficient in the medium to long term and that transplantation should be viewed as a tool of last resort. We argue that there has been too much focus on transplanting fast-growing branching corals, which in general naturally recruit well but tend to survive transplantation and relocation relatively poorly, to create short-term increases in live coral cover, at the

expense of slow-growing massive corals, which generally survive transplantation well but often recruit slowly. In those cases where transplantation is justified, we advocate that a reversed stance, which focuses on early addition of slowly recruiting massive species to the recovering community, rather than a short-term and sometimes short-lived increase in coral cover, may be more appropriate in many cases. Edwards, AJ, Clark, S, Zahir, H, Rajasuriya, A, Naseer, A and Rubens, J. (2001) Coral bleaching and mortality on artificial and natural reefs in Maldives in 1998, sea surface temperature anomalies and initial recovery. Marine Pollution Bulletin 42, 7-15. The bleaching and subsequent mortality of branching and massive corals on artificial and natural reefs in the central atolls of Maldives in 1998 are examined with respect to sea surface temperature (SST) anomalies. SST normally peaks in April-May in Maldives. The UK Meteorological Office's Global sea-Ice and SST data set version 2.3b shows that in 1998 monthly mean SST was 1.2-4 S.D. above the 1950-1999 average during the warmest months (March-June), with the greatest anomaly in May of +2.1°C. Bleaching was first reported in mid-April and was severe from late April to mid-May with some recovery evident by late-May. At least 98% of branching corals (Acroporidae, Pocilloporidae) on artificial structures deployed on a reef flat in 1990 died whereas the majority of massive corals (Poritidae, Faviidae, Agariciidae) survived the bleaching. The pre-bleaching coral community on the artificial reefs in 1994 was 95% branching corals and 5% massives (n=1589); the post-bleaching community was 3% branching corals and 97% massives (n=248). Significant reductions in live coral cover were seen at all natural reefs surveyed in the central atolls, with average live coral cover decreasing from about 42% to 2%, a 20-fold reduction from pre-bleaching levels. A survey of recruitment of juvenile corals to the artificial structures 10 months after the bleaching event showed that 67% of recruits (>0.5 cm diameter) were acroporids and pocilloporids and 33% were from massive families (n=202) compared to 94% and 6%, respectively, in 1990-1994 (n=3136). Similar post-bleaching dominance of recruitment by branching corals was seen on nearby natural reef (78% acroporids and pocilloporids; 22% massives). A linear regression of April mean monthly SST against year was highly significant (p<0.001) and suggests a rise of 0.16°C per decade. If this trend continues, by 2030 mean April SST in the central atolls will normally exceed the anomaly level at which corals appear there are susceptible to mass bleaching. Einbinder, S, Perelberg, A, Ben-Shaprut, O, Foucart, MH and Shashar, N. (2006) Effects of artificial reefs on fish grazing in their vicinity: Evidence from algae presentation experiments. Marine Environmental Research 61, 110-119. Artificial reefs have been suggested as a tool for conservation and restoration of marine habitats. However, the relationships between coral reef habitats and man-made structures are poorly understood. We experimentally tested whether artificial reefs change grazing patterns in their surrounding environment. We exposed heaps of the macroalgae, Ulva lactuca, to natural grazing, at various distances from three artificial reefs. Results suggest that artificial reefs change grazing patterns in the neighboring area. In all the locations examined grazing was 2–3 times higher near the artificial reefs than in control sites (p<0.05). We suggest that herbivorous fishes are attracted to the artificial reefs, creating a zone of increased grazing. Therefore,

while planning deployment of such artificial reefs it is necessary to consider their overall influence on their natural surroundings, in order to maintain the natural community trophic dynamics. Elvidge, CD, Dietz, JB, Berkelmans, R, Andréfouët, S, Skirving, W, Strong, AE and Tuttle, BT. (2004) Satellite observation of Keppel Islands (Great Barrier Reef) 2002 coral bleaching using IKONOS data. Coral Reefs 23, 123-132. An examination of IKONOS satellite imagery of the Keppel Islands (Great Barrier Reef) acquired before and during a coral bleaching event indicates that severe bleaching of reefs can be detected as an increase in brightness in the band 1 (blue) and band 2 (green) IKONOS spectral bands (4-m resolution). The bleaching was not detected in band 3 (red), band 4 (near-infrared), or in the 1-m panchromatic band data. A total of 0.74 km2 of bleached coral was identified, with detection occurring in waters as deep as 15 m. The procedure requires that one of the scenes be radiometrically normalized to match the reference scene prior to image differencing. A relative radiometric normalization was used in this case because variable cloud cover present in the image acquired during the bleaching event prevented reliable modeling of atmospheric effects. The success at coral bleaching detection at Keppel Islands represents both a ‘‘best-case’’ and a ‘‘cloud-challenged’’ scenario. It was a best-case scenario in that coral cover was extensive (70–90% live coral cover, mostly acroporids) and the bleaching level was extreme (92–95% of coral cover white bleached). It was a cloud-challenged scenario in terms of having extensive and highly variable cloud cover present in the image acquired during the bleaching event. Color difference images reveal extensive areas of bleached coral at sites away from our study area, indicating that this platform and methodology may be a valuable tool for mapping high coral cover areas during bleaching events. Additional studies and technique refinements would be required to test the detection limits of bleaching with IKONOS imagery or to develop a spectrally based bleaching detection index. Epstein, N, Bak, RPM and Rinkevich, B. (2001) Strategies for gardening denuded coral reef areas: The applicability of using different types of coral material for reef restoration. Restoration Ecology 9, 432-442. Recreational and other human activities degrade coral reefs worldwide to a point where efficient restoration techniques are needed. Here we tested several strategies for gardening denuded reefs. The gardening concept consists of in situ or ex situ mariculture of coral recruits, followed by their transplantation into degraded reef sites. In situ nurseries were established in Eilat's (Northern Red Sea) shallow waters, sheltering three types of coral materials taken from the branching species Stylophora pistillata (small colonies, branch fragments, and spat) that were monitored for up to two years. Pruning more than 10% of donor colonies' branches increased mortality, and surviving colonies displayed reduced reproductive activity. Maricultured isolated branches, however, exceeded donor colony life span and reproductive activity and added 0.5-45% skeletal mass per year. Forty-four percent of the small colonies survived after 1.5-year mariculture, revealing average yearly growth of 75 ± 32%. Three months ex situ maintenance of coral spat (sexual recruits) prior to the in situ nursery phase increased survivorship. Within the next 1.5 years, they developed into colonies of 3-4 cm diameter. Nursery periods of 2 years, 4-5 years, and more than >5 years have been estimated for small colonies, spat, and isolated branches,

respectively. These and other results, including the possible use of nubbins (minute fragments the size of a single or few polyps), are discussed, revealing benefits and drawbacks for each material. In situ coral mariculture is an improved practice to the common but potentially harmful protocol of direct coral transplantation. It is suggested that reef gardening may be used as a key management tool in conservation and restoration of denuded reef areas. The gardening concept may be applicable for coral reefs worldwide through site-specific considerations and the use of different local coral species. Epstein, N, Bak, RPM and Rinkevich, B. (2003) Applying forest restoration principles to coral reef rehabilitation. Aquatic Conservation: Marine and Freshwater Ecosystems 13, 387-395. 1. Forest restoration through silviculture (gardening) programs revives productivity, biodiversity, and stability. As in silviculture approaches, the coral ‘gardening’ strategy is based on a two-step protocol. 2. The first step deals with the establishment of in situ and/or ex situ coral nurseries in which corals are farmed (originating from two types of source material: asexual [ramets, nubbins], and sexual [planula larvae, spat] recruits). 3. The second is the reef rehabilitation step, where maricultured colonies are transplanted into degraded sites. 4. We compare here the rationale of forest restoration to coral reef ecosystem restoration by evaluating major key criteria. As in silviculture programs, a sustainable mariculture operation that focuses on the prime structural component of the reef (‘gardening’ with corals) may promote the persistence of threatened coral populations, as well as that of other reef taxa, thus maintaining genetic diversity. In chronically degrading reef sites this may facilitate a halt in biodiversity depletion. 5. Within the current theoretical framework of ecosystem restoration, the recovery of biodiversity indices is considered a core element since a rich species diversity provides higher ecosystem resilience to disturbances. 6. The gardening measure may also be implemented worldwide, eliminating the need to extract existing colonies for transplantation operations. At degraded reef sites, the coral gardening strategy can assist in managing human and non-human stakeholders’ requirements as is done in forest management. Epstein, N, Vermeij, MJA, Bak, RPM and Rinkevich, B. (2004) Alleviating impacts of anthropogenic activities by traditional conservation measures: Can a small reef reserve be sustainably managed? Biological Conservation. Zoning schemes that are becoming an important management tool in large marine reserves, are difficult to implement in small reef areas. At the 3.4 km long reef of Eilat (Red Sea), a small (ca. 350 m of coastline) enclosure strategy has been enforced since 1992, while the remaining reef was left open to intense human activities. Here we have investigated for 2.5 y three populations of the branching coral Stylophora pistillata (3605 colonies) in a locality within the enclosed area (site NR) and in two areas open to the public, by tossing random quadrats at the shallow lagoonar zone (0.5–1.5 m depth). In the two open sites we found significantly higher levels of colony breakage (14–34% vs. 4–9% in the enclosed site), lower partial mortality levels of colonies (in the first 1.5 y; 7–9% vs. 23–30% at NR), higher recruitment (up to 3.0 vs. up to 0.9 colonies/m 2), 50% reduction in coral life span (10 vs. 20 y) and an

estimated extinction period of 9–10 y for new cohorts as compared to >20 y in the enclosed site. Average colony size and maximal colony size were about half in the open sites. Live coverage fluctuated widely in all sites but was 3 times higher in the enclosed area (1.0–3.0% vs. 0.3–1.1%). Log-transformed size frequency distributions revealed, at the open sites, a shift from small towards medium-size classes and at the enclosed site, a shift from larger to medium size classes. We conclude that the enclosure of a limited core zone, although improved some ecological parameters, was not sufficient to compensate for stress imposed by anthropogenic activities. It is suggested to employ active restoration approaches, such as the ‘‘gardening concept’’, as supplementary management tools. Fabricius, KE, Wild, C, Wolanski, E and Abele, D. (2003) Effects of transparent exopolymer particles and muddy terrigenous sediments on the survival of hard coral recruits. Estuarine, Coastal and Shelf Science 57, 613-621. Sedimentation is a major cause of mortality in scleractinian coral recruits. In this study, we compared the effects of muddy coastal sediments, with and without enrichment by ‘marine snow’, on the survivorship of recruits of the hard coral Acropora willisae. Transparent exopolymer particles (TEP) were measured as characteristic components of marine snow using a staining method (Passow & Alldredge, Limnol. Oceangr. 40 (1995) 1326). Four-week-old recruits were exposed to: (1) muddy coastal sediments; (2) TEP; (3) TEP-enriched muddy coastal sediments; and (4) unfiltered sea water, for 43 h in aerated flow chambers. Thirty-three percent (± SE) of coral recruits died after 43-h exposure to TEP-enriched muddy coastal sediments (~14 mgcm-2 sediments enriched with 3.8 ± 0.2 µg cm-2 gum xanthan equivalents (GX) TEP). In contrast, no or minimal mortality was observed in the other three treatments. Mortality increased to >80% when the amount of deposited TEP was almost tripled (10.9 ± 1.3 µg cm-2 GX) and sediment increased by 50%. Thus, coral recruits survived short-term exposure to low levels of TEP and low levels of muddy sediments, but sediments enriched with TEP at concentrations recorded at some of the inshore stations proved to be detrimental. Concentrations of TEP were measured in the central Great Barrier Reef (latitude 16-18°S) in summer, the season of coral spawning and recruitment. Within <10km off the coast, TEP concentrations were high (mean=291 ± 49 SE µgGXl-1, range=152–791 µgGXl-1). Concentrations declined with increasing distance from the coast, and averaged 83 (±26 SE) µgGXl-1 around oceanic reefs >40km off the coast. Our study suggests that both sediment composition and short-term (43 h) sediment deposition affect survival of coral juveniles, which has implications for the capacity of inshore reefs to be recolonised by corals to recover from acute disturbance events. Fabricius, KE, Mieog, JC, Colin, PL, Idip, D and van Oppen, MJ. (2004) Identity and diversity of coral endosymbionts (zooxanthellae) from three Palauan reefs with contrasting bleaching, temperature and shading histories. Molecular Ecology 13, 2445-2458. The potential of corals to associate with more temperature-tolerant strains of algae (zooxanthellae, Symbiodinium) can have important implications for the future of coral reefs in an era of global climate change. In this study, the genetic identity and diversity of zooxanthellae was investigated at three reefs with contrasting histories of bleaching mortality, water temperature and shading, in the Republic of Palau

(Micronesia). Single-stranded conformation polymorphism and sequence analysis of the ribosomal DNA internal transcribed spacer (ITS)1 region was used for genotyping. A chronically warm but partly shaded coral reef in a marine lake that is hydrographically well connected to the surrounding waters harboured only two single-stranded conformation polymorphism profiles (i.e. zooxanthella communities). It consisted only of Symbiodinium D in all 13 nonporitid species and two Porites species investigated, with the remaining five Porites harbouring C*. Despite the high temperature in this lake (> 0.5 degrees above ambient), this reef did not suffer coral mortality during the (1998) bleaching event, however, no bleaching-sensitive coral families and genera occur in the coral community. This setting contrasts strongly with two other reefs with generally lower temperatures, in which 10 and 12 zooxanthella communities with moderate to low proportions of clade D zooxanthellae were found. The data indicate that whole coral assemblages, when growing in elevated seawater temperatures and at reduced irradiance, can be composed of colonies associated with the more thermo-tolerant clade D zooxanthellae. Future increases in seawater temperature might, therefore, result in an increasing prevalence of Symbiodinium phylotype D in scleractinian corals, possibly associated with a loss of diversity in both zooxanthellae and corals. Fagoonee, I, Wilson, HB, Hassell, MP and Turner, J. (1999) The dynamics of zooxanthellae populations: A long-term study in the field. Science 283, 843-845. Coral bleaching characterized by the expulsion of symbiotic algae (zooxanthellae) is an increasing problem worldwide. Global warming has been implicated as one cause, but the phenomenon cannot be fully comprehended without an understanding of the variability of zooxanthellae populations in field conditions. Results from a 6-year field study are presented, providing evidence of density regulation but also of large variability in the zooxanthellae population with regular episodes of very low densities. These bleaching events are likely to be part of a constant variability in zooxanthellae density caused by environmental fluctuations superimposed on a strong seasonal cycle in abundance. Fautin, DG and Buddemeier, RW. (2004) Adaptive bleaching: A general phenomenon. Hydrobiologia 530/531, 459-467. Laboratory and field data bearing on the adaptive bleaching hypothesis (ABH) are largely consistent with it; no data of which we are aware refute it. We generalize the ABH in light of these data and observations. The population of zooxanthellae within an organism is dynamic, the diversity of zooxanthellae is both surprising and difficult to ascertain, and field experiments demonstrate both turn-over in zooxanthella types and habitat-holobiont correlations. Dynamic change in symbiont communities, and the idea of an equilibrium or optimal community that matches the environment at a particular place and time, are concepts that underlie or emerge from much of the recent literature. The mechanism we proposed to explain responses to acute bleaching appears to operate continuously, thereby enabling the host-symbiont holobiont to track even subtle environmental changes and respond promptly to them. These findings enhance the potential importance of the ABH in the outcomes of acute bleaching, which can (1) accelerate this process of holobiont change, and (2) change the set of possible trajectories for how symbiont communities might recover.

Fiksel, J. (2006) Sustainability and resilience: Toward a systems approach. Sustainability: Science, Practice and Policy. A comprehensive systems approach is essential for effective decision making with regard to global sustainability, since industrial, social, and ecological systems are closely linked. Despite efforts to reduce unsustainability, global resource consumption continues to grow. There is an urgent need for a better understanding of the dynamic, adaptive behavior of complex systems and their resilience in the face of disruptions, recognizing that steady-state sustainability models are simplistic. However, assessing the broad impacts of policy and technology choices is a formidable challenge, as exemplified in life-cycle analysis of the implications of alternative energy and mobility technologies. A number of research groups are using dynamic modeling techniques, including biocomplexity, system dynamics, and thermodynamic analysis, to investigate the impacts on ecological and human systems of major shifts such as climate change and the associated policy and technology responses. These techniques can yield at least a partial understanding of dynamic system behavior, enabling a more integrated approach to systems analysis, beneficial intervention, and improvement of resilience. Recommendations are provided for continued research to achieve progress in the dynamic modeling and sustainable management of complex systems. Fine, M and Loya, Y. (2002) Endolithic algae: An alternative source of photoassimilates during coral bleaching. Proceedings of the Royal Society of London B Biological Sciences 269, 1205-1210. Recent reports of worldwide coral bleaching events leading to devastating coral mortality have caused alarm among scientists and resource managers. Differential survival of coral species through bleaching events has been widely documented. We suggest that among the possible factors contributing to survival of coral species during such events are endolithic algae harboured in their skeleton, providing an alternative source of energy. We studied the dynamics of photosynthetic pigment concentrations and biomass of endoliths in the skeleton of the encrusting coral Oculina patagonica throughout a bleaching event. During repeated summer bleaching events these endolithic algae receive increased photosynthetically active radiation, increase markedly in biomass, and produce increasing amounts of photoassimilates, which are translocated to the coral. Chlorophyll concentrations and biomass of endoliths were 4.6 ± 1.57 and 1570 ± 427 µg cm-2 respectively, in skeletons of relatively healthy colonies (0–40% bleaching) but up to 14.8 ± 2.5 and 4036 ± 764 µg cm-2 endolith chlorophyll and biomass respectively, in skeletons of bleached colonies (greater than 40% bleaching). The translocation dynamics of 14C-labelled photoassimilates from the endoliths to bleached coral tissue showed significantly higher 14C activity of the endoliths harboured within the skeletons of bleached corals than that of the endoliths in non-bleached corals. This alternative source of energy may be vital for the survivorship of O. patagonica, allowing gradual recruitment of zooxanthellae and subsequent recovery during the following winter.

Fine, M, Steindler, L and Loya, Y. (2004) Endolithic algae photoacclimate to increased irradiance during coral bleaching. Marine and Freshwater Research 55, 115-121. The photoacclimation of endolithic algae (of the genus Ostreobium) inhabiting the skeleton of the Mediterranean coral Oculina patagonica during a bleaching event was examined. Pulse amplitude modulated (PAM) chlorophyll fluorescence techniques in situ were used to assess the photosynthetic efficiency of endolithic algae in the coral skeleton and the symbiotic dinoflagellates (zooxanthellae) in the coral tissue. Relative photosynthetic electron transport rates (ETRs) of the endolithic algae under bleached areas of the colony were significantly higher than those of endolithic algae from a healthy section of the colony and those of zooxanthellae isolated from the same section. Endolithic algae under healthy parts of the colony demonstrated an ETR[max] of 16.5% that of zooxanthellae from tissue in the same section whereas endolithic algae under bleached sections showed ETR[max] values that were 39% of those found for healthy zooxanthellae. The study demonstrates that endolithic algae undergo photoacclimation with increased irradiance reaching the skeleton. As PAM fluorometry has become a major tool for assessing levels of stress and bleaching in corals, the importance of considering the contribution of the endolithic algae to the overall chlorophyll fluorescence measured is highlighted. Finelli, CM, Helmuth, BST, Pentcheff, ND and Wethey, DS. (2006) Water flow influences oxygen transport and photosynthetic efficiency in corals. Coral Reefs 25, 47-57. Recent studies indicate that the incidence and persistence of damage from coral reef bleaching are often highest in areas of restricted water motion, and that resistance to and recovery from bleaching is increased by enhanced water motion. We examined the hypothesis that water motion increases the efflux of oxygen from coral tissue thereby reducing oxidative stress on the photosynthetic apparatus of endosymbiotic zooxanthellae. We experimentally exposed colonies of Montastrea annularis and Agaricia agaricites to manipulations of water flow, light intensity, and oxygen concentration in the field using a novel mini-flume. We measured photosynthetic efficiency using a pulse amplitude modulated fluorometer to test the short-term response of corals to our manipulations. Under normal oxygen concentrations, A. agaricites showed a significant 8% increase in photosynthetic efficiency from 0.238 (± 0.032) in still water to 0.256 (± 0.037) in 15 cm s-1 flow, while M. annularis exhibited no detectable change. Under high-ambient oxygen concentrations, the observed effect of flow on A. agaricites was reversed: photosynthetic efficiencies showed a significant 11% decrease from 0.236 (± 0.056) in still water to 0.211 (± 0.048) in 15 cm s-1 flow. These results support the hypothesis that water motion helps to remove oxygen from coral tissues during periods of maximal photosynthesis. Flow mitigation of oxidative stress may at least partially explain the increased incidence and severity of coral bleaching in low flow areas and observations of enhanced recovery in high-flow areas.

Fitt, WK, Spero, HJ, Halas, J, White, MW and Porter, JW. (1993) Recovery of the coral Montastrea annularis in the Florida Keys after the 1987 Caribbean "bleaching event". Coral Reefs 12, 57-64. Many reef-building corals and other cnidarians lost photosynthetic pigments and symbiotic algae (zooxanthellae) during the coral bleaching event in the Caribbean in 1987. The Florida Reef Tract included some of the first documented cases, with widespread bleaching of the massive coral Montastrea annularis beginning in late August. Phototransects at Carysfort Reef showed discoloration of >90% of colonies of this species in March 1988 compared to 0% in July 1986; however no mortality was observed between 1986 and 1988. Samples of corals collected in February and June 1988 had zooxanthellae densities ranging from 0.1 in the most lightly colored corals, to 1.6 x 106 cells/cm2 in the darker corals. Minimum densities increased to 0.5 x 106 cells/cm2 by August 1989. Chlorophyll-a content of zooxanthellae and zooxanthellar mitotic indices were significantly higher in corals with lower densities ofzooxanthellae, suggesting that zooxanthellae at low densities may be more nutrient sufficient than those in unbleached corals. Ash-free dry weight of coral tissue was positively correlated with zooxanthellae density at all sample times and was significantly lower in June 1988 compared to August 1989. Proteins and lipids per cm 2 were significantly higher in August 1989 than in February or June, 1988. Although recovery of zooxanthellae density and coral pigmentation to normal levels may occur in less than one year, regrowth of tissue biomass and energy stores lost during the period of low symbiont densities may take significantly longer. Fitt, WK, McFarland, FK, Warner, ME and Chilcoat, GC. (2000). Seasonal patterns of tissue biomass and densities of symbiotic dinoflagellates in reef corals and relation to coral bleaching. Limnology and Oceanography 45, 677-685. Tissue biomass (ash-free dry weight) and symbiotic dinoflagellates (density, chlorophyll a cell-1 or cm-2 of coral surface area) of five species of reef-building corals were monitored seasonally for up to 4 yr at three different depths in the Bahamas. The lowest values of all tissue biomass and algal symbiont parameters occurred during the late summer-fall sample periods. In contrast, the highest densities and pigment content of symbionts usually occurred during the winter, whereas tissue biomass peaked most often in the spring, the time lag implying a functional relationship between these variables. Corals living in shallow water often (but not always) had higher levels of all parameters measured compared to deeper corals, except chlorophyll a content, which usually displayed the opposite trend. The results show that corals from all depths exhibited bleaching (loss of symbiotic dinoflagellates and/or their pigments) every year, regardless of whether they appeared white, tan, or mottled to the human eye. We speculate that these patterns are driven by seasonal changes in light and temperature on algal and animal physiology. Furthermore, we hypothesize that all tropical reef-building corals, world-wide, exhibit similar predictable cycles in their tissue biomass and symbiotic algae.

Fitt, WK, Brown BE, Warner, ME and Dunne, RP. (2001) Coral bleaching: Interpretation of thermal tolerance limits and thermal thresholds in tropical corals. Coral Reefs 20, 51-65. It should be clear that the upper temperature limit for life cannot be accurately defined' (Schmidt-Nielsen 1996). The thermal physiology of zooxanthellate reef corals is reviewed in this paper in the context of organismal and biochemical responses occurring during coral bleaching, with emphasis on methods of detection and interpretation of animal and algal symbiont stress. Coral bleaching, as presently defined in the literature, is a highly subjective term used to describe a variety of conditions pertaining to low symbiont densities in the coral-algal complex, including response to thermal stress. Three general types of high-temperature bleaching are defined: physiological bleaching, which may or may not include higher-than-normal temperature responses; algal-stress bleaching, involving dysfunction of symbiotic algae at high light and/or high temperatures; and animal-stress bleaching, where coral cells containing symbiotic algae are shed from the gastrodermal layer of cells. Since none of these methods of bleaching is mutually exclusive, a combination of intrusive and non-intrusive techniques is necessary to determine which mechanisms of symbiont loss are occurring. While quantification of symbiont densities, algal pigments, and coral tissue biomass provide unambiguous evidence of bleaching severity, measurements of physiological and biochemical degradation offer additional correlative evidence of temperature stress. Pulse-amplitude-modulated (PAM) fluorometry has emerged as an easy and relatively inexpensive non-invasive technique for monitoring symbiotic algal function both in situ and in the laboratory, when proper assumptions and interpretations are made. The roles of global warming, water quality, acclimation/adaptation processes, and relation to coral disease and reef heterogeneity are also discussed. A thorough understanding of the organismal responses occurring during bleaching will help explain changes in coral populations and in the coral reef community, and perhaps assist in predicting the future of reef corals and coral reefs during the next century of global climate change. Floros, CD, Samways, MJ and Armstrong, B. (2004) Taxonomic patterns of bleaching within a South African coral assemblage. Biodiversity and Conservation 13, 1175-1194. In 1998, the Indian Ocean coral reefs suffered a severe and extensive mass bleaching event. The thermal tolerances of corals were exceeded and their photosynthetic symbionts (zooxanthellae) lost. Mortalities of up to 90% were recorded on the reefs of Seychelles, Maldives, Kenya and Tanzania. South African coral reefs were among the few that largely escaped the 1998 mass bleaching event, but may be threatened in the future if global warming increases. This study assessed the extent of coral bleaching and partial recovery at Sodwana Bay, South Africa during 2000 and 2001. Bleaching levels in this study varied over the course of a year, which suggested that seasonally varying parameters such as sea temperature were the most likely cause of bleaching. Bleaching levels were highest at the shallowest site. However, these bleaching levels were very low in comparison with those of reefs elsewhere in the Indian Ocean. The greater volume of water over the relatively deeper reefs of Sodwana Bay may have protected the reefs from severe bleaching. Field measurements on the three reefs indicated that, although the reefs at Sodwana Bay are still healthy, bleaching increased from <1% in 1998 to 5–10% in 2002.

Bleaching occurred in 26 coral genera. The Alcyonacea were highly susceptible to bleaching, especially Sarcophyton sp. Among the hard corals, Montipora spp. Were the species most susceptible to bleaching. The sensitivity of these genera to early and slight increases in temperature suggests that they can forewarn of a possible greater bleaching event. In contrast, the coral genera Turbinaria and Stylophora were most resistant to bleaching. Folke, C. (2002) Social-ecological resilience and behavioural responses. In Biel, A, Hansson, B and Mårtensson, M. ‘Individual and structural determinants of environmental practice.’ Ashgate Aldershot (UK), 226-242. Abstract not available. Fong, P and Glynn, PW. (2000) A regional model to predict coral population dynamics in response to El Niño-Southern Oscillation. Ecological Applications 10, 842-854. The objective of this paper was to modify an existing size-structured population model for the coral Gardineroseris planulata developed from field measurements at the Uva Island reef, Panama, for use at the regional scale (Tropical Eastern Pacific). The modified model can be used to aid in prediction of changes in coral population dynamics in response to ENSO events of different frequency and magnitude. The earlier model incorporates coral growth, predation by the sea star Acanthaster planci, and mortality associated with El Niño–Southern Oscillation (ENSO) events. The regional model was modified from the earlier local version by extending the coral growth–temperature relationship to include the lower temperatures observed in areas of strong seasonal upwelling. In addition, ENSO-associated mortality was modified by incorporating a dynamic function that relates partial mortality of corals in different size classes to the rate of temperature change during an ENSO event. This function is based on data from three reef areas (Uva Island and the Pearl Islands, Panama, and the Galápagos Islands, Ecuador), which represent a regional gradient from an area characterized by relatively high annual temperature with low variability to areas with lower annual temperature subject to seasonal upwelling. Model predictions of coral population size structure in different areas agreed with field observations. Predictions for coral recovery after an ENSO event were most accurate for the Uva Island population and least accurate for the Galapágos Islands population. We also tested the effect of various ENSO frequencies on the size structure of the coral population. Increasing ENSO frequency in the upwelling area resulted in an increase in the proportion of small colonies, while the same increase in ENSO frequency in the thermally stable reef area had little effect on the population. Sensitivity analysis suggested that more accurate estimates of growth of the smallest and largest colonies in upwelling areas as well as predation on intermediate-size colonies are important to improve predictive capability. Fortes, MD, Djohani, R and Kahn, B. (2002) Introducing the Regional Action Plan (RAP) to strengthen a resilient network of effective Marine Protected Areas in Southeast Asia: 2002-2012. World Commission on Protected Areas: Sanur (Indonesia), 23 pp. No abstract available.

Fox, HE, Pet, JS, Dahuri, R and Caldwell, RL. (2003) Recovery in rubble fields: Long-term impacts of blast fishing. Marine Pollution Bulletin. This paper presents initial results from a study of factors that inhibit or enhance hard coral recovery in rubble fields created by blast fishing in Komodo National Park and Bunaken National Park, Indonesia. Within nine sites monitored since 1998, there was no significant natural recovery. Levels of potential source coral larvae were assessed with settlement tiles in the rubble fields and in nearby high coral cover sites. Rubble movement was measured and shown to be detrimental to small scleractinians, especially in high current areas. In shallow water (2–6 m deep), rubble is often overgrown by soft corals and corallimorpharians, which inhibit hard coral survival. There is increased scleractinian recruitment in quadrats cleared of soft coral, and Acropora nubbins transplanted into soft coral fields suffer greater mortality than those transplanted above the soft coral canopy. Gaining an understanding of the prognosis for coral recovery is essential not only in order to assess the long-term impacts of blast fishing, but also to improve management decisions about protection of intact reefs and potential restoration of damaged areas. Fox, HE, Mous, PJ, Pet, JS, Muljadi, AH and Caldwell, RL. (2005) Experimental assessment of coral reef rehabilitation following blast fishing. Conservation Biology 19, 98-107. Illegal fishing with explosives has damaged coral reefs throughout Southeast Asia. In addition to killing fish and other organisms, the blasts shatter coral skeletons, leaving fields of broken rubble that shift in the current, abrading or burying new coral recruits, and thereby slowing or preventing reef recovery. Successful restoration and rehabilitation efforts can contribute to coral reef conservation. We used field experiments to assess the effectiveness of different low-cost methods for coral reef rehabilitation in Komodo National Park (KNP), Indonesia. Our experiments were conducted at three different spatial scales. At a scale of 1 × 1 m plots, we tested three different rehabilitation methods: rock piles, cement slabs, and netting pinned to the rubble. Significantly more corals per square meter grew on rocks, followed by cement, netting, and untreated rubble, although many plots were scattered by strong water current or buried by rubble after 2.5 years. To test the benefits of the most successful treatment, rocks, at more realistic scales, we established 10 × 10 m plots of rock piles at each of our nine sites in 2000. Three years after installation, coverage by hard corals on the rocks continued to increase, although rehabilitation in high current areas remained the most difficult. In 2002 rehabilitation efforts in KNP were increased over 6000 m2 to test four rock pile designs at each of four rubble field sites. Assuming that there is an adequate larval supply, using rocks for simple, low-budget, large-scale rehabilitation appears to be a viable option for restoring the structural foundation of damaged reefs.

Franklin, DJ, Cedrés, CMM and Hoegh-Guldberg, O. (2006) Increased mortality and photoinhibition in the symbiotic dinoflagellates of the Indo–Pacific coral Stylophora pistillata (Esper) after summer bleaching. Marine Biology 149, 633–642. 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 midsummer, 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 2.5·106 to 0.8·106 dinoflagellates cm2 coral tissue), and those dinoflagellates that remained after summer became photoinhibited more readily (dark-adapted FV : Fm 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. Freire, FFM. (2001) The application of Geographic Information System to the coral reef of southern Okinawa. Proceedings of the 22nd Asian Confernce on Remote Sensing: Singapore, 110-115. Abstract not available. See www.crisp.nus.edu.sg/~acrs2001/pdf/194freire.pdf Fujiwara, S and Omori, M. (2004) Coral reef restoration. In Japanese Ministry of the Environment and Japanese Coral Reef Society (eds.) ‘Coral reefs of Japan’: Tokyo (Japan), 138-145. Abstract not available. See www.coremoc.go.jp/english/pub/coralreefjapan/0501_restoration_techniques.pdf Gardner, TA, Côte, IM, Gill, JA, Grant, A and Watkinson, AR. (2003) Long-term region-wide declines in Caribbean corals. Science 301, 958-960. We report a massive region-wide decline of corals across the entire Caribbean basin, with the average hard coral cover on reefs being reduced by 80%, from about 50% to 10% cover, in three decades. Our meta-analysis shows that patterns of change in coral cover are variable across time periods but largely consistent across subregions, suggesting that local causes have operated with some degree of synchrony on a region-wide scale. Although the rate of coral loss has slowed in the past decade compared to the 1980s, significant declines are persisting. The ability of Caribbean

coral reefs to cope with future local and global environmental change may be irretrievably compromised. Garren, M, Walsh, SM, Caccone, A and Knowlton, N. (2006) Patterns of association between Symbiodinium and members of the Montastraea annularis species complex on spatial scales ranging from within colonies to between geographic regions. Coral Reefs 25, 503–512. Patterns of associations between coral colonies and the major clades of zooxanthellae can vary across scales ranging from individual colonies to widely separated geographic regions. This is exemplified in this study of the Montastraea annularis species complex from six sites on the Mesoamerican Reef, Belize and nine sites in the Bocas del Toro archipelago, Panama. Restriction fragment length polymorphism (RFLP) analysis of small subunit ribosomal DNA (SSU rDNA) was used to identify the zooxanthellae. In Belize (M. annularis), Symbiodinium B (79% of the colonies), Symbiodinium A, and Symbiodinium C were observed. In Panama (primarily M. franksi, but also M. annularis and M. faveolata), there was greater diversity and evenness with Symbiodinium A, B, C, C’ (a new symbiont) and D all being common in at least some host/habitat combinations. Non-metric multidimensional scaling ordinations showed that distribution patterns of symbionts across sites are best explained by enclosure (relative influence of open ocean vs. coastal water) and total suspended solids. Because members of clade D are known to be temperature resistant and Symbiodinium C’ was found in environments characterized by high sedimentation, these Panamanian reefs may have importance from a management perspective as reservoirs of corals better able to tolerate human impacts. Gates, RD, Baghdasarian, G and Muscatine, L. (1992) Temperature Stress Causes Host Cell Detachment in Symbiotic Cnidarians: Implications for Coral Bleaching. Biological Bulletin 182, 324-332. During the past decade, acute and chronic bleaching of tropical reef corals has occurred with increasing frequency and scale. Bleaching, i.e., the loss of pigment and the decrease in population density of symbiotic dinoflagellates (zooxanthellae), is often correlated with an increase or decrease in sea surface temperature. Because little is known of the cellular events concomitant with thermal bleaching, we have investigated the mechanism of release of zooxanthellae by the tropical sea anemone Aiptasia pulchella and the reef coral Pocillopora damicornis in response to cold and heat stress. Both species released intact host endoderm cells containing zooxanthellae. The majority of the released host cells were viable, but they soon disintegrated in the seawater leaving behind isolated zooxanthellae. The detachment and release of intact host cells suggests that thermal stress causes host cell adhesion dysfunction in these cnidarians. Knowledge of the cellular entity released by the host during bleaching provides insight into both the underlying release mechanism and the way in which natural environmental stresses evoke a bleaching response.

Gates, RD, Bil, KY and Muscatine, L. (1999) The influence of an anthozoan ‘‘host factor’’ on the physiology of a symbiotic dinoflagellate. Journal of Experimental Marine Biology and Ecology 232, 241–259. Anthozoan host factors are components of host animal homogenates which elicit the release of newly fixed carbon by symbiotic dinoflagellates in vitro. The host factor active component of an homogenate of the reef coral Pocillopora damicornis has recently been isolated and identified as a mixture of free amino acids. A synthetic host factor (SHF) based on the measured free amino acid pool of P. damicornis elicits an identical response in symbiotic dinoflagellates in vitro as does the crude host factor. SHF was used as a tool with which to investigate the impact of anthozoan host factor on other aspects of symbiotic dinoflagellate metabolism. Here we demonstrate that dinoflagellates isolated from the tropical sea anemone Aiptasia pulchella (Carlgren) incubated in SHF, in addition to releasing fixed carbon, also exhibit higher photosynthetic carbon fixation, heterotrophic carbon fixation, photosynthetic oxygen production, respiration rates, and cell specific concentrations of chlorophyll a as compared to dinoflagellates incubated in sea water in vitro. Thus, the performance of symbiotic dinoflagellates in vitro appears to be sustained in an amino acid rich environment, but compromised in sea water. This observation raises the possibility that in nature symbiotic dinoflagellates released from anthozoan hosts may survive in amino acid rich environments such as fish guts and ‘‘intermediate hosts’’. Gates, RD and Edmunds, PJ. (1999) The physiological mechanisms of acclimatization in tropical coral reefs. American Zoologist 39, 30-41. The ability of scleractinian corals to survive changes that are predicted in the global environment over the next century will lie in their physiological mechanisms of acclimatization. Corals display rapid modifications in behavior, morphology and physiology enabling them to photoacclimate to changing light conditions, a scenario that demonstrates considerable biological flexibility. Here we argue that the acclimatization mechanisms in corals are fundamentally similar to those exhibited by other invertebrate taxa. We discuss protein metabolism as a mechanism underlying acclimatization responses in reef corals, and explore the relationship between protein turnover, metabolic rate, growth rate, and acclimatization capacity. Our preliminary analyses suggest that corals with low growth rates ([micro]Ca/mgN/h) and high metabolic rates ([micro][O.sub.2]/cm2/hr), such as the massive species, acclimatize more effectively than those with high growth rates and low metabolic rates, a feature that is characteristic of branching species. We conclude that studies of protein turnover, combined with temporally relevant investigations into the dynamic aspects of coral dinoflagellate symbioses will provide considerable insight into why corals exhibit such a high level of variation in response to the same environmental challenge. Furthermore, a more detailed understanding of acclimatization mechanisms is essential if we are to predict how a coral assemblage will respond to present and future environmental challenges.

Gibson, J, McField, M and Wells, S. (1998) Coral reef management in Belize: An approach through Integrated Coastal Zone Management. Ocean and Coastal Management 39, 229-244. Belize has one of the most extensive reef ecosystems in the Western Hemisphere, comprising one of the largest barrier reefs in the world, three atolls and a complex network of inshore reefs. Until recently, the main impacts were probably from natural events such as hurricanes. However, anthropogenic threats such as sedimentation, agrochemical run-off, coastal development. tourism and overfishing are now of concern. To limit these impacts, Belize is taking the approach of integrated coastal zone management. The programme is building on the existing legislative framework and involves the development of an appropriate institutional structure to co-ordinate management activities in the coastal zone. A Coastal Zone Management Plan is being prepared, which will include many measures that will directly benefit the reefs: a zoning scheme for the coastal zone, incorporating protected areas; legislation and policy guidelines: research and monitoring programmes; education and public awareness campaigns; measures for community participation; and a financial sustainability mechanism. Glassom, D, Zakai, D and Chadwick-Furman, NE. (2004) Coral recruitment: A spatio-tempral analysis along the coastline of Eilat, northern Red Sea. Marine Biology 144, 641-651. Recruitment rates of stony corals to artificial substrates were monitored for 2 years at 20 sites along the coast of Eilat, northern Red Sea, to compare with those recorded at other coral reef locations and to assess variation in recruitment at several spatial scales. Coral recruitment was low compared to that observed on the Great Barrier Reef in Australia, but was similar to levels reported from other high-latitude reef locations. Pocilloporids were the most abundant coral recruits in all seasons. Recruitment was twofold higher during the first year than during the second year of study. There was considerable spatial variability, with the largest proportion of variance, apart from the error term, attributable to differences between sites, at a scale of 102 m. Spearman’s ranked correlation showed consistency in spatial patterns of recruitment of pocilloporid corals between years, but not of acroporid corals. During spring, when only the brooding pocilloporid coral Stylophora pistillata reproduces at this locality, most coral recruitment occurred at central and southern sites adjacent to well-developed coral reefs. During summer, recruitment patterns varied significantly between years, with wide variation in the recruitment of broadcasting acroporid corals at northern sites located distant from coral reefs. Settlement was low at all sites during autumn and winter. This work is the first detailed analysis of coral recruitment patterns in the Red Sea, and contributes to the understanding of the spatial and temporal scales of variation in this important reef process.

Glassom, D, Celliers, L and Schleyer, MH. (2006) Coral recruitment patterns at Sodwana Bay, South Africa. Coral Reefs 25, 485–492. Recruitment is widely acknowledged as one of the most important processes in the maintenance of coral reef systems, particularly in their recovery and replenishment following disturbances. In this study variation in coral recruitment was monitored for 3 years at Sodwana Bay, South Africa, located in the Greater St Lucia Wetland Park (GSLWP), a world heritage site. Due to the latitude (27–28°S) and physical conditions, this area is considered marginal for coral reef growth. Recruitment of corals to ceramic tiles peaked between March and May each year, with broadcast spawners dominating plates in March and brooders more abundant in May. Pocilloporid corals were the most abundant during all periods except March 2001, when acroporids comprised 72% of the total number of spat. Total recruitment was the highest during this period, with a mean of 13.2 ± 15.73 (mean ± SD) corals tile-1. Recruitment rates varied widely between and within different reefs monitored. Patterns of variation were inconsistent between seasons, but some reefs had persistently low rates of settlement. Glick, P. (1999) Coral reefs and climate change: Last straw for a threatened ecosystem. National Wildlife Federation: Washington DC (USA), 8 pp. Little can compare to the natural beauty and rich biological diversity of the world’s coral reefs. From the Great Barrier Reef ecosystem “down under” to the series of reefs off the Florida Keys, these colorful “rainforests of the sea” are home to thousands of marine species — one of every four ocean species known. Their structures help protect coasts from erosion and storm damage. And they support a fishing and tourism industry worth hundreds of billions of dollars globally. Unfortunately, these glorious ecosystems are under siege by human activities. Increasingly, the world’s reefs are being ravaged by dredging and coral collecting, poisoned by pollution and cyanide, and depleted by over-fishing. Furthermore, the evidence is mounting that global warming — the result of extensive use of fossil fuels and destruction of the world’s forests — is dealing these already weakened ecosystems a serious blow. Without a concerted global effort to protect and restore coral reefs from all of these threats, we stand to lose more than just an important resource — we will lose one of nature’s crown jewels and a fundamental part of our oceans’ ecological vitality. The National Wildlife Federation (NWF) has produced this report to help you learn more about the threats to coral reef ecosystems and what all of us can do to save these natural treasures, for the benefit of people and wildlife alike! Global Environmental Facility Targeted Research Group (2005) Coral reef targeted research and capacity building for management. GEF: Washington DC (USA), 16 pp. The Coral Reef Targeted Research and Capacity Building program has been established to address fundamental information gaps in our understanding of coral reef ecosystems, so that management options and policy interventions can be strengthened globally. For the first time in history, this project will join the collective effort of many of the World’s leading coral reef scientists to coordinate research efforts and address key outstanding questions about the health of coral reefs. The

program is being developed in phases over 15 years, and through focused and systematic research, is working to effectively support management and policy and to better integrate resulting information with other disciplines, such as economics and law. The program will also enhance the capacity of researchers, students and managers within developing countries, so that a global network can effectively share the most up-to-date research to benefit regional, national and local management actions and policy. Golbuu, Y, Victor, S, Penland, L, Idip Jr, D, Emaurois, C, Okaji, K, Yukihira, H, Iwase, A, and van Woesik, R. (2007) Palau’s coral reefs show differential habitat recovery following the 1998-bleaching event. Coral Reefs 26, 319–332. Documenting successional dynamics of coral communities following large-scale bleaching events is necessary to predict coral population responses to global climate change. In 1998, high sea surface temperatures and low cloud cover in the western Pacific Ocean caused high coral mortality on the outer exposed reefs of Palau (Micronesia), while coral mortality in sheltered bays was low. Recovery was examined from 2001 to 2005 at 13 sites stratified by habitat (outer reefs, patch reefs and bays) and depth (3 and 10 m). Two hypotheses were tested: (1) rates of change of coral cover vary in accordance with habitat, and (2) recovery rates depend on recruitment. Coral cover increased most in the sheltered bays, despite a low recruitment rate, suggesting that recovery in bays was primarily a consequence of remnant regrowth. Recruitment densities were consistently high on the wave-exposed reefs, particularly the western slopes, where recovery was attributed to both recruitment and regrowth of remnants. Recovery was initially more rapid at 10 m than 3 m on outer reefs, but in 2004, recovery rates were similar at both depths. Rapid recovery was possible because Palau’s coral reefs were buffered by remnant survival and recruitment from the less impacted habitats. Gorbunov, MY, Kolber, ZS, Lesser, MP and Falkowski, PG. (2001) Photosynthesis and Photoprotection in Symbiotic Corals. Limnology and Oceanography 46, 75-85. In zooxanthellate corals, excess excitation energy can be dissipated as heat (nonphotochemical quenching), thereby providing protection against oxidative damage by supraoptimal light in shallow reefs. To identify and quantify the photoprotective mechanisms, we studied the diel variability of chlorophyll fluorescence yields and photosynthetic parameters in situ in corals, using moored and SCUBA-based fast-repetition-rate fluorometers. The results reveal that nonphotochemical quenching is triggered prior to saturation of photosynthetic electron transport by downregulation of the reaction centers of Photosystem I1 (PSII). This process dissipates up to 80% of the excitation energy. On a sunny day in shallow waters, the daily integrated flux of photons absorbed, and subsequently dissipated as heat, is -4 times that used for photosynthesis. Fluorescence quenching is further accompanied by a slight reduction in the functional absorption cross section for PSII that results from thermal dissipation of excitation energy in the light-harvesting antennae. These two processes are highly dynamic and adjust to irradiance changes on timescales consistent with the passage of clouds across the sky. Under supraoptimal irradiance, however, up to 30% of PSII reaction centers become photoinhibited, and these are repaired only after several hours of low

irradiance. In shallow corals, between 10% and 20% of the reactions centers are chronically photoinhibited and appear to remain permanently nonfuncitonal throughout the year. Our results establish, for the first time, the suite of biophysical mechanisms that optimize photosynthesis while simultaneously providing photoprotection in symbiotic corals in situ. Goreau, TJ. (1998) Coral recovery from bleaching in Alphonse and Bijoutier, Seychelles. Coral Reef Alliance: Cambridge (USA). Mortality of corals in the Alphonse group following the 1998 bleaching event was somewhat less than in the high islands of the Mahe group. Overall mortality of corals on fore reef slopes was around 90%in shallow water and around 70-80% on vertical slopes, in comparison to values around 95 and 99% in the inner islands. Although mortality of Acroporas was nearly total, several small surviving colonies were found at a range of depths including very shallow seagrasses. In contrast with the high islands, many partially surviving Pocillopora verrucosa colonies were found. The major survivors were Helioporacoerula (rare in the inner islands), Porites head species (probably lutea), Porites cylindrica, Porites nigrescens, Astreoporamyriophthalma, Hydnophora microconos, and Favid corals. Coral diversity was lower than in the high islands and no Diploastreas or Gonioporas were seen in the Alphonse group, although these were found at every site in the high islands. Survival of corals was very high, roughly 80%, inside the Alphonse lagoon, also higher than equivalent habitats in the Inner Islands. Although the tiny human population is insufficient to cause eutrophication, evidence of high natural nutrient sources was seen. Strong upwelling is the cause of significant algae abundance in deep waters, while the lagoons at Alphonse and Saint Francois (but not Bijoutier, which lacks a lagoon) have green water which affects surrounding reefs and promotes algae growth, apparently as the result of recycling of nutrients from sea grass decomposition. Areas not affected by such sources, such as Bijoutier, are predominantly overgrown by encrusting pink calcareous algae, creating the surfaces needed for new coral settlement. Goreau, TJ, McClanahan, TR, Hayes, R and Strong, A. (2000) Conservation of coral reefs after the 1998 global bleaching event. Conservation Biology 14, 5-15. Abstract not available. Goreau, TJ. (2003) Predicting coral bleaching by HotSpot analysis. Coral Reef Alliance: Cambridge (USA). GCRA researchers originated and developed the Coral Bleaching Hotspot method that has been used to successfully forecast all major coral bleaching events since 1990, and to hind cast all major bleaching events since 1982. Typically we are able to predict when and where bleaching will take place and how bad it will be using this method, often before the first signs are visible in the field, allowing field researchers to be warned to catch the early stages of such events.

Goreau, TJ, Cervino, JM and Pollina, R. (2004) Increased zooxanthellae numbers and mitotic index in electrically stimulated corals. Symbiosis 37, 107-120. Zooxanthella densities, mitotic indices, and chlorophyll were measured in six major reef-building coral genera growing on electrically stimulated "Biorock" reefs in Indonesia, and compared with genetically identical corals growing adjacent to them. Corals on Biorock reefs had generally higher densities of zooxanthellae and higher rates of symbiotic algal division, but generally lower chlorophyll per zooxanthella. These patterns are coincident with higher coral skeletal growth rates, better developed branching morphology, and higher rates of resistance to environmental stresses, including elevated temperatures, nutrients, and sediments. Zooxanthellae play a critical role in providing alkalinity for coral calcification by withdrawing carbon dioxide for photosynthesis, and energy for proton and calcium pumping. Our results support the view that Biorock corals are healthier than normal corals because alkalinity for calcification is provided by elevated pH caused by electrolysis of seawater. Goreau, TJ. (2005) Global warming and coral reefs. Open Democracy: London (UK), 3 pp. Coral reefs are the most sensitive of all ecosystems to global warming, pollution, and new diseases. They will be first to go as a result of climate change. As the most important resources for fisheries, tourism, shore protection, and marine biodiversity for more than a hundred countries, this will be a huge disaster. Goulet, TL and Coffroth, MA. (2003) Stability of an octocoral-algal symbiosis over time and space. Marine Ecology Progress Series 250, 117-124. In symbiosis, 2 taxonomically different organisms co-exist, each pursuing their own agenda and yet, they are linked in one entity. A mutualistic symbiosis may break up if it is no longer beneficial to either one of the partners. Changing needs over time or changing environmental conditions may prompt symbiont switching. For example, corals may survive elevated temperatures by switching their algal symbionts. If switching occurs, the new combination of host and symbiont genotypes may perform better. Conversely, the partners may be fixed for life, with the degree to which the mutualism responds to changing selection pressures dictated by the existing partners. Understanding the genotypic dynamics of a mutualism is important for predicting the potential resilience of a mutualism over time and in the face of environmental perturbations. Although mutualisms tend to be characterized at the species level or higher, host-symbiont dynamics is an individual-level question, requiring individual-level analysis. We used multilocus DNA fingerprinting to examine long-term temporal and spatial symbiont change in the mutualism between the octocoral Plexaura kuna and its algal symbionts (zooxanthellae). We monitored zooxanthella genotypes within a colony for up to 10 yr, among P. kuna clonemates, across different habitats and in colonies transplanted to novel environments. In all instances, the prominent zooxanthella genotype within a P. kuna colony remained unchanged although zooxanthella genotypes varied among genetically distinct P. kuna colonies. Such tremendous temporal and spatial stability may occur in other

coral hosts, influencing the reaction and survival of mutualisms during environmental change. Goulet, TL. (2006) Most corals may not change their symbionts. Marine Ecology Progress Series 321, 1-7. Many corals (stony corals and octocorals) rely on their symbiotic algae (zooxanthellae)

for survival. Under stress, zooxanthellae are expelled, resulting in coral bleaching. The hypothesis that corals may survive climate change by exchanging algal types is shown here to be potentially applicable only to a minority of corals. Data on 442 coral species from 43 studies reveal that only a few coral species may be able to change their symbionts. The ability to change symbionts seems to be linked to whether a coral species can host multiple zooxanthella clades, either at different depths on the same reef, on different reefs or at different geographic locations, or concurrently within the same colony. The combined data set shows that only 23% of coral species host multiple zooxanthella clades. Most coral species (77%) exhibit fidelity to a narrow subset of a single zooxanthella clade, some even to specific algal genotypes within a clade. These specific algal genotypes in coral species hosting a single algal clade do not change over time. Furthermore, no algal change occurs when a coral colony is either transplanted to different environments, or subjected to stressors such as disease or increased temperatures. For the majority of corals, therefore, algal switching does not appear to occur. These coral species will survive only if the existing host–symbiont combination withstands the changing conditions. If climate warming continues, coral reefs may undergo a change in biodiversity such that only a subset of symbiotic corals may persist. Graham, NAJ, Wilson, SK, Jennings, S, Polunin, NVC, Bijoux, JP and Robinson, J. (2006) Dynamic fragility of oceanic coral reef ecosystems. Proceedings of the National Academy of Sciences 103, 8425-8429. As one of the most diverse and productive ecosystems known, and one of the first ecosystems to exhibit major climate-warming impacts (coral bleaching), coral reefs have drawn much scientific attention to what may prove to be their Achilles heel, the thermal sensitivity of reef-building corals. Here we show that climate change-driven loss of live coral, and ultimately structural complexity, in the Seychelles results in local extinctions, substantial reductions in species richness, reduced taxonomic

distinctness, and a loss of species within key functional groups of reef fish. The importance of deteriorating physical structure to these patterns demonstrates the longer-term impacts of bleaching on reefs and raises questions over the potential for recovery. We suggest that isolated reef systems may be more susceptible to climate change, despite escaping many of the stressors impacting continental reefs Graham, NAJ, McClanahan, TR, Letourneur, Y and Galzin, R. (2007) Anthropogenic stressors, inter-specific competition and ENSO effects on a Mauritian coral reef. Environmental Biology of Fishes 78, 57-69. Much of the western Indian Ocean suffered widespread loss of live coral in 1998 and interest is now focussed on the indirect effects of this coral loss on other components of the ecosystem, in particular fishes. However, it is just as important to identify changes in fish assemblages at locations that did not suffer coral mortality to

understand local versus regional drivers. We surveyed benthic and fish communities on a reef flat in Mauritius five times between 1994 and 2005. The design allowed for comparison through time, along the coast and between inshore and offshore reef locations. The benthic community demonstrates a clear trend along the coast, likely in response to a dredged water ski lane, but little change through time. Branching Acropora colonies dominate much of the live coral and best explain patterns in the fish assemblage (P < 0.01). Few changes in overall fish species richness through time were identified, and observed changes were within fishery target families rather than species reliant on live coral. Departure from expected levels of taxonomic distinctness suggests degradation in the community associated with the dredged ski lane. Non-metric multi-dimensional scaling of the fish assemblage demonstrates a similar pattern to that seen in the benthos; greater differences along the coast (Global R = 0.34) than through time (Global R = 0.17) and no trend between reef positions. SIMPER analysis identified two species of Stegastes as the main drivers of trends in the MDS plot and the most dominant of these, S. lividus, appears to be reducing species richness of the remaining fish community. The study highlights Mauritius as a regional refugia of thermally-sensitive corals and specialised fish, suggesting a need for careful management. Graham, NAJ, Wilson, SK, Jennings, S, Polunin, NVC, Robinson, J, Bijoux, JP and Daw, TM. (2007) Lag Effects in the Impacts of Mass Coral Bleaching on Coral Reef Fish, Fisheries, and Ecosystems. Conservation Biology. Recent episodes of coral bleaching have led to wide-scale loss of reef corals and raised concerns over the effectiveness of existing conservation and management efforts. The 1998 bleaching event was most severe in the western Indian Ocean, where coral declined by up to 90% in some locations. Using fisheries-independent data, we assessed the long-term impacts of this event on fishery target species in the Seychelles, the overall size structure of the fish assemblage, and the effectiveness of two marine protected areas (MPAs) in protecting fish communities. The biomass of fished species above the size retained in fish traps changed little between 1994 and 2005, indicating no current effect on fishery yields. Biomass remained higher in MPAs, indicating they were effective in protecting fish stocks. Nevertheless, the size structure of the fish communities, as described with size-spectra analysis, changed in both fished areas and MPAs, with a decline in smaller fish (<30 cm) and an increase in larger fish (>45 cm). We believe this represents a time-lag response to a reduction in reef structural complexity brought about because fishes are being lost through natural mortality and fishing, and are not being replaced by juveniles. This effect is expected to be greater in terms of fisheries productivity and, because congruent patterns are observed for herbivores, suggests that MPAs do not offer coral reefs long-term resilience to bleaching events. Corallivores and planktivores declined strikingly in abundance, particularly in MPAs, and this decline was associated with a similar pattern of decline in their preferred corals. We suggest that climate-mediated disturbances, such as coral bleaching, be at the fore of conservation planning for coral reefs.

Grandcourt, EM and Cesar, HSJ. (2003) The bio-economic impact of mass coral mortality on the coastal reef fisheries of the Seychelles. Fisheries Research 60, 539-550. Data collected through a stratified catch and effort survey were used to assess the impact of the 1998 mass coral bleaching event on socio-economic and biological indicators for the coastal reef fisheries of the Seychelles. There was a significant reduction in the abundance index and monthly yields per square kilometre for representatives of the family Siganidae following 1998. However, this was not associated with the bleaching event and conformed with the declining trend prior to the impact. Abundance indices and yields per square kilometre did not change significantly for the primary target families of the handline fisheries (Lutjanidae, Serranidae, Lethrinidae and Carangidae). Declining trends in abundance indices and yields for Octopodidae reversed after 1998, although the phenomena could not be independently linked to coral bleaching. Whilst critical resource based management issues are identified for the demersal handline fishery, the results suggest that there were no negative short-term bio-economic impacts on Seychelles coastal reef fisheries associated with mass coral mortality. Great Barrier Reef Marine Park Authority. (1998) Bleaching – The great unknown. Reef Research 8.

Recent mass bleaching of corals in the southern hemisphere has received much attention and speculation as to its occurrence. Simply put: nobody really knows for sure. The phenomenon of coral bleaching was noted as early as the late 1920s during the Great Barrier Reef Expedition. However, mass bleaching events have only been recorded since the late 1970s and became a more closely studied event in the 1980s. Some of the factors that are thought to cause the bleaching are elevated sea temperatures, exposure to excessive irradiance and lowered salinity. However, when corals approach their upper thermal limits, even small additional doses of ultra-violet light or other sunlight spectra can cause them to bleach. In some parts of the world coral bleaching has occurred every three to four years since the late ´70s. The subsequent research has led to increased knowledge about the event but much is still unsubstantiated.

Great Barrier Reef Marine Park Authority. (2006) Climate change and the Great Barrier Reef. GBRMPA: Townsville (Australia), http://www.gbrmpa.gov.au/corp_site/info_services/science/climate_change/

There is now overwhelming scientific and political consensus that climate change is occurring as a result of human activity, and that it is one of the biggest threats to reefs worldwide. The major predictions associated with climate change, due largely to the continuing increases in the concentrations and effects of greenhouse gases, include:

• Increases in average sea surface temperature; • Increases in average sea level; • Increased El Nino events; • Changes to rainfall patterns;

• Changes to ocean circulation; and • Increased concentrations of CO2 in the ocean resulting in altered ocean

chemistry.

The Australian Greenhouse Office report Climate Change Risk and Vulnerability predicts that increasing temperature and sea level rise are two of the biggest climate change threats to Australia in the next 50 years. These climate change predictions may have serious consequences for the Great Barrier Reef.

Griffin, SP and Bhagooli, R. (2004) Measuring antioxidant potential in corals using the FRAP assay. Journal of Experimental Marine Biology and Ecology 302, 201-211. In this paper, we standardized a method for determining antioxidant potential in corals. This was determined using a simple, reproducible and inexpensive method: the ferric reducing/antioxidant potential (FRAP) assay. This procedure involves the reduction of FeIII-TPTZ to a blue colored FeII-TPTZ by biological antioxidants and chemical reductants, some of which might have no antioxidant activity in a sample. The FRAP assay compares the change in absorbance at 600 nm of a sample compared with the change in absorbance of a known standard (FeSO4·7H2O) to determine antioxidant levels. This assay was used to determine changes in antioxidant potential in the corals Pocillopora damicornis and Pocillopora meandrina exposed to different temperatures (28, 29, 30 and 31 °C) for 3 h. Corals were also incubated at 31 °C for time intervals of a 0.5, 1 and 3 h. Antioxidant potential in the coral host increased with temperature and time, as indicated by FRAP values, compared to control samples at ambient sea surface temperatures (26.5–27 °C). Lower FRAP values could be a response to lower production of reactive oxygen species (ROS) or the result of an increase in ROS that react with the antioxidants. Because of the complex interactions within cells, one test is normally not enough to understand precisely what is going on within the cell. Rather, a broad array of tests is required to determine the different cellular parameters that are occurring within a biological system. To our knowledge, this is the first time that FRAP has been used to determine antioxidant status in a marine organism. The FRAP technique can potentially be a useful and inexpensive tool for marine biologists engaged in ecotoxicological studies. Grimsditch, GD and Salm, RV. (2006) Coral reef resilience and resistance to bleaching. IUCN and The Nature Conservancy: Gland (Switzerland), 53 pp. Coral reefs are vital ecosystems, providing a source of income, food and coastal protection for millions of people; and recent studies have shown that coral reef goods and services provide an annual net benefit of US$30 billion to economies worldwide (Cesar et al, 2003). Coral reefs are composed mainly of reef-building corals: colonial animals (polyps) that live symbiotically with the singlecelled microalgae (zooxanthellae) in their body tissue and secrete a calcium carbonate skeleton. Coral reefs are formed by hundreds of thousands of these polyps and are found in warm, shallow, clear, low-nutrient tropical and sub-tropical waters, with optimum temperatures of 25-29ºC, although they exist in ranges from 18ºC (Florida) to 33ºC (Persian Gulf) (Buddemeier and Wilkinson, 1994). They are incredibly diverse, covering only 0.2% of the ocean’s floor but containing 25% of its species and they

are often dubbed the ‘tropical rainforests of the oceans’ (Roberts, 2003). Unfortunately, coral reefs are also among the most vulnerable ecosystems in the world. Disturbances such as bleaching, fishing, pollution, waste disposal, coastal development, sedimentation, SCUBA diving, anchor damage, predator outbreaks, invasive species and epidemic diseases have all acted synergistically to degrade coral reef health and resilience. Today, an estimated 20% of coral reefs worldwide have been destroyed, while 24% are in imminent danger and a further 26% are under longer term danger of collapse (Wilkinson, 2004). This paper synthesises much of the current scientific knowledge on coral reef resistance and resilience to bleaching, a possible major effect of climate change. Following a brief overview of coral bleaching and what is meant by resistance and resilience, the paper highlights a variety of resistance and resilience factors and identifies some gaps in knowledge. It continues by providing an overview of some of the tools and strategies we can use to enhance coral reef resilience. Finally, it reviews current initiatives working on coral reef resilience and also identifying some possible future opportunities for research into the issue. Grottoli, AG, Rodrigues, LJ and Palardy, JE. (2006) Heterotrophic plasticity and resilience in bleached corals. Nature 440, 1186-1189. Mass coral bleaching events caused by elevated seawater temperatures have resulted in extensive coral mortality throughout the tropics over the past few decades. With continued global warming, bleaching events are predicted to increase in frequency and severity, causing up to 60% coral mortality globally within the next few decades. Although some corals are able to recover and to survive bleaching, the mechanisms underlying such resilience are poorly understood. Here we show that the coral host has a significant role in recovery and resilience. Bleached and recovering Montipora capitata (branching) corals met more than 100% of their daily metabolic energy requirements by markedly increasing their feeding rates and CHAR (per cent contribution of heterotrophically acquired carbon to daily animal respiration), whereas Porites compressa (branching) and Porites lobata (mounding) corals did not. These findings suggest that coral species with high-CHAR capability during bleaching and recovery, irrespective of morphology, will be more resilient to bleaching events over the long term, could become the dominant coral species on reefs, and may help to safeguard affected reefs from potential local and global extinction. Guinotte, JM, Buddemeier, RW and Kleypas, JA. (2003) Future coral reef habitat marginality: Temporal and spatial effects of climate change in the Pacific basin. Coral Reefs 22, 551-558. Marginal reef habitats are regarded as regions where coral reefs and coral communities reflect the effects of steady-state or long-term average environmental limitations. We used classifications based on this concept with predicted time-variant conditions of future climate to develop a scenario for the evolution of future marginality. Model results based on a conservative scenario of atmospheric CO2 increase were used to examine changes in sea surface temperature and aragonite saturation state over the Pacific Ocean basin until 2069. Results of the projections indicated that essentially all reef locations are likely to become marginal with respect to aragonite saturation state. Significant areas, including some with the highest

biodiversity, are expected to experience high-temperature regimes that may be marginal, and additional areas will enter the borderline high temperature range that have experienced significant ENSO-related bleaching in the recent past. The positive effects of warming in areas that are presently marginal in terms of low temperature were limited. Conditions of the late 21st century do not lie outside the ranges in which present-day marginal reef systems occur. Adaptive and acclimative capabilities of organisms and communities will be critical in determining the future of coral reef ecosystems. Gunderson, L. (2006) Ecology: A Different Route to Recovery for Coral Reefs. Current Biology 17, 27-28. Worldwide, many coral reef ecosystems have undergone regime shifts, changing from domination by coral to domination by algae. New work indicates that the return path is surprisingly different fromthe forward one. Guzman, HM and Cortés, J. (2007) Reef recovery 20 years after the 1982–1983 El Niño massive mortality. Marine Biology 151, 401-411. For over 20 years the El Niño-Southern Oscillation (ENSO) has caused damage to the coral reefs of the eastern PaciWc and other regions. In the mid-1980s scientists estimated that coral cover was reduced by 50–100% in several countries across the region. Almost 20 years (2002) after the 1982–1983 event, we assessed the recovery of the virtually destroyed reefs at Cocos Island (Costa Rica), previously evaluated in 1987 and reported to have less than 4% live coral cover. We observed up to fivefold increase in live coral cover which varied among reefs surveyed in 1987 and 2002. Most new recruits and adults belonged to the main reef building species from pre-1982 ENSO, Porites lobata, suggesting that a disturbance as outstanding as El Niño was not sufficient to change the role or composition of the dominant species, contrary to phase shifts reported for the Caribbean. During the 1990s, new species were observed growing on the reefs. Notably, Leptoseris scabra, considered to be rare in the entire Pacific, was commonly found in the area. Recovery may have begun with the sexual and asexual recruits of the few surviving colonies of P. lobata and Pavona spp. and with long distance transport of larvae from remote reefs. We found an overall 23% live coral cover by 2002 and with one reef above 58% indicating that Cocos Island coral reefs are recovering. Hallock, P. (2005) Global change and modern coral reefs: New opportunities to understand shallow-water carbonate depositional cycles. Sedimentary Geology 175, 19-33. Human activities are impacting coral reefs physically, biologically, and chemically. Nutrification, sedimentation, chemical pollution, and overfishing are significant local threats that are occurring worldwide. Ozone depletion and global warming are triggering mass coral-bleaching events; corals under temperature stress lose the ability to synthesize protective sunscreens and become more sensitive to sunlight. Photo-oxidative stress also reduces fitness, rendering reef-building organisms more susceptible to emerging diseases. Increasing concentration of atmospheric CO2 has already reduced CaCO3 saturation in surface waters by more than 10%. Doubling of atmospheric CO2 concentration over pre-industrial concentration in the 21st century

may reduce carbonate production in tropical shallow marine environments by as much as 80%. As shallow-water reefs decline worldwide, opportunities abound for researchers to expand understanding of carbonate depositional systems. Coordinated studies of carbonate geochemistry with photozoan physiology and calcification, particularly in cool subtropical-transition zones between photozoan-reef and heterotrophic carbonate-ramp communities, will contribute to understanding of carbonate sedimentation under environmental change, both in the future and in the geologic record. Cyanobacteria are becoming increasingly prominent on declining reefs, as these microbes can tolerate strong solar radiation, higher temperatures, and abundant nutrients. The responses of reef-dwelling cyanobacteria to environmental parameters associated with global change are prime topics for further research, with both ecological and geological implications. Halpern, BS, Selkoe, KA, Micheli, F and Kappel, CV. (2007) Evaluating and Ranking the Vulnerability of Global Marine Ecosystems to Anthropogenic Threats. Conservation Biology. Marine ecosystems are threatened by a suite of anthropogenic stressors. Mitigating multiple threats is a daunting task, particularly when funding constraints limit the number of threats that can be addressed. Threats are typically assessed and prioritized via expert opinion workshops that often leave no record of the rationale for decisions, making it difficult to update recommendations with new information. We devised a transparent, repeatable, and modifiable method for collecting expert opinion that describes and documents how threats affect marine ecosystems. Experts were asked to assess the functional impact, scale, and frequency of a threat to an ecosystem; the resistance and recovery time of an ecosystem to a threat; and the certainty of these estimates. To quantify impacts of 38 distinct anthropogenic threats on 23 marine ecosystems, we surveyed 135 experts from 19 different countries. Survey results showed that all ecosystems are threatened by at least nine threats and that nine ecosystems are threatened by >90% of existing threats. The greatest threats (highest impact scores)were increasing sea temperature, demersal destructive fishing, and point-source organic pollution. Rocky reef, coral reef, hard-shelf, mangrove, and offshore epipelagic ecosystems were identified as the most threatened. These general results, however, may be partly influenced by the specific expertise and geography of respondents, and should be interpreted with caution. This approach to threat analysis can identify the greatest threats (globally or locally), most widespread threats, most (or least) sensitive ecosystems, most (or least) threatened ecosystems, and other metrics of conservation value. Additionally, it can be easily modified, updated as new data become available, and scaled to local or regional settings, which would facilitate informed and transparent conservation priority setting. Hansen, LJ, Biringer, JL and Hoffman, JR. (2003) Buying time: A user’s manual for building resistance and resilience to climate change in natural systems. WWF Climate Change Program: Berlin (Germany), 244 pp. As we stand at the beginning of the new millennium, the threats to nature and protected areas are unprecedented. While some progress has been made and strategies such as protected areas have been successful in preserving biodiversity in some places, new threats are arising. None of these threats is as great as global

climate change and none will have such large implications for the way natural resource managers plan and implement conservation strategies. While global climate change is seemingly difficult to understand and plan for, planning is essential, as the conservation approaches of the past may not work in an ever-changing warmer world. New strategies, led by deep cuts in the heat-trapping gases that causes climate change, predominately carbon dioxide from the burning of fossil fuels, may at least buy some time for ecosystems to adapt in the years and decades ahead. However, if CO2 emissions are not reduced quickly and deeply, some of those treasured ecosystems will not survive. Climate change is happening now and nature is experiencing its impacts first. Whether one looks at coral reefs, mangroves, arctic areas or montane regions, climate change poses a complex and bewildering array of problems for ecosystems. The key question is, what can be done—in addition to the rapid reduction of CO2 emissions now—to increase the resiliency of these ecosystems to climate change? WWF compiled “Buying Time: A User’s Manual for Building Resistance and Resilience to Climate Change in Natural Systems” for natural resource managers who are ready to confront the impacts of climate change. While far from comprehensive, this manual brings together assessments and potential initial adaptation strategies for various biomes. Written by experts, the manual addresses all of the major biomes with practical ideas of how to begin increasing the resiliency of ecosystems and plan our protected areas in response to the threat of climate change. Some of these strategies are in line with the conservation strategies we have been working on for years—reducing fragmentation, building corridors, reducing threats, and increasing resiliency in general. Natural resource managers in the field must begin planning to buy time for these biomes while the world works to switch from coal to clean power, a key strategy to reduce the causes of climate change. Looking at the models and gathering empirical evidence, WWF is recommending that natural resource managers start now to build climate change adaptation strategies into their preservation philosophies and plans. This manual is a first step to assist managers in doing so. While testing and implementing these new conservation strategies, managers should also communicate the threat that climate change poses to their biome to local, regional and national authorities. Resource managers have an important role to play in the climate change debate by using examples of changes seen in their regions as indicators of the need for rapid and deep cuts in CO2 emissions. Harding, S, Solandt, J-L, Walker, RCJ, Walker, D, Taylor, J, Haycock, S, Davis, MT and Raines, PS. (2003) Reef Check data reveals rapid recovery from coral bleaching in the Mamanucas, Fiji. Silliman Journal 44, 80-98. Twenty two fringing reef sites within the Mamanuca Islands, western Fiji were surveyed during 2001 and 2002, using Reef Check methods. A mean increase of 14.3% in hard coral cover was recorded over the 12-month period. This increase in hard coral cover suggests a significant recovery of scleractinian coral colonies that were originally impacted by the 2000 mass bleaching episode in the South Pacific. The event was reported to have caused >80% coral mortality in the southern and eastern regions of Fiji. Between 2001 and 2002 the coral reefs of the Mamanucas progressed from .poor. to .fair. in accordance with the Association of South East Asian Nations (ASEAN) system for describing the health of coral reefs. Our results

also show that trained non-specialist volunteers undertaking marine surveys such as Reef Check can competently collect simple, yet important quantitative data regarding the physical health of coral reefs. Harithsa, S, Raghukumar, C and Dalal, SG. (2005) Stress response of two coral species in the Kavaratti atoll of the Lakshadweep Archipelago, India. Coral Reefs. Frequent occurrences of coral bleaching and the ensuing damage to coral reefs have generated interest in documenting stress responses that precede bleaching. The objective of this study was to assess and compare physiological changes in healthy, semi-bleached andtotally bleached colonies of two coral species, Porites lutea and Acropora formosa, during a natural bleaching event in the Lakshadweep Archipelago in the Arabian Sea to determine the traits that will be useful in the diagnosis of coral health. In April 2002, three ‘‘health conditions’’ were observed as ‘‘appearing healthy,’’ ‘‘semi-bleached’’ and ‘‘bleached’’ specimens for two dominant and co-occurring coral species in these islands. Changes in the pigment composition, zooxanthellae density (ZD), mitotic index (MI) of zooxanthellae, RNA/DNA ratios and protein profile in the two coral species showing different levels of bleaching in the field were compared to address the hypothesis of no difference in health condition between species and bleaching status. The loss in chlorophyll (chl) a, chl c and ZD in the transitional stage of semi-bleaching in the branched coral A. formosa was 80, 75 and 80%, respectively. The losses were much less in the massive coral P. lutea, being 20, 50 and 25%, respectively. The decrease in zooxanthellar density and chl a was accompanied by an increased MI of zooxanthellae and RNA/DNA ratios in both the species. There was an increase in accumulation of lipofuscin granules in partially bleached P. lutea tissue, which is an indication of cellular senescence. Multivariate statistical analyses showed that colonies of P. lutea ranked in different health conditions differed significantly in chl a, chl c, ZD, RNA/DNA ratios, and protein concentrations, whereas in A. formosa chl a, chl c, chl a/c, phaeopigments and MI contributed to the variance between health conditions. Harley, CDG, Hughes, AR, Hultgren, KM, Miner, BG, Sorte, CJB, Thornber, CS, Rodriguez, LF, Tomanek, L and Williams, SL. (2006) The impacts of climate change in coastal marine systems. Ecology Letters 9, 228-241. Anthropogenically induced global climate change has profound implications for marine ecosystems and the economic and social systems that depend upon them. The relationship between temperature and individual performance is reasonably well understood, and much climate-related research has focused on potential shifts in distribution and abundance driven directly by temperature. However, recent work has revealed that both abiotic changes and biological responses in the ocean will be substantially more complex. For example, changes in ocean chemistry may be more important than changes in temperature for the performance and survival of many organisms. Ocean circulation, which drives larval transport, will also change, with important consequences for population dynamics. Furthermore, climatic impacts on one or a few ‘leverage species’ may result in sweeping community-level changes. Finally, synergistic effects between climate and other anthropogenic variables, particularly fishing pressure, will likely exacerbate climate-induced changes. Efforts to manage and conserve living marine systems in the face of climate change will require

improvements to the existing predictive framework. Key directions for future research include identifying key demographic transitions that influence population dynamics, predicting changes in the community-level impacts of ecologically dominant species, incorporating populations’ ability to evolve (adapt), and understanding the scales over which climate will change and living systems will respond. Helmuth, BST, Timmerman, BEH and Sebens, KP. (1997) Interplay of host morphology and symbiont microhabitat in coral aggregations. Marine Biology 130, 1-10. The Belizean reef coral Agaricia tenuifolia Dana forms aggregations in which rows of thin, upright blades line up behind each other. On average, the spacing between blades increases with depth and hence with decreasing ambient irradiance. We designed and built a small, inexpensive light meter and used it to quantify the effect of branch spacing on light levels within colonies at varying distances from branch tips. Concurrently, we measured photosynthetic pigment concentrations and population densities of symbiotic dinoflagellates (zooxanthellae) extracted from coral branches of colonies with tight (≤3 cm) vs wide (≥6 cm) branch spacing, collected at 15 to 17 m and from colonies with tight branch spacing collected at 1 to 2 m. Light levels decreased significantly with tighter branch spacing and with distance from the branch tips. Total cellular pigment concentrations (chlorophylls a, c2 and peridinin) as well as chlorophyll a:c2 and chlorophyll a:peridinin ratios all increased significantly with distance from the branch tip, indicating very localized differences in photoacclimation within individual branches. Zooxanthellae from colonies with widely-spaced branches displayed significantly lower chlorophyll a:c2 and chlorophyll a:peridinin ratios, and were present at significantly higher population densities than those from colonies with tightly-spaced branches collected at the same depth (15 m). Tightly-spaced colonies collected from shallow environments (1 to 2 m) displayed pigment ratios similar to those from widely-spaced colonies from deeper water (15 m), but maintained zooxanthellae populations at levels similar to those in tightly-branched colonies from deeper water. Thus, variation in colony morphology (branch spacing and distance from branch tip) can affect symbiont physiology in a manner comparable to an increase of over 15 m of water depth. These results show that a host's morphology can strongly determine the microhabitat of its symbionts over very small spatial scales, and that zooxanthellae can in turn display steep gradients in concordance with these altered physical conditions. Helmuth, B, Kingsolver, JG and Carrington, E. (2005) Biophysics, Physiological ecology and climate change: Does mechanism matter? Annual Review of Physiology 67, 177–201. Recent meta-analyses have shown that the effects of climate change are detectable and significant in their magnitude, but these studies have emphasized the utility of looking for large-scale patterns without necessarily understanding the mechanisms underlying these changes. Using a series of case studies, we explore the potential pitfalls when one fails to incorporate aspects of physiological performance when predicting the consequences of climate change on biotic communities. We argue that by considering the mechanistic details of physiological performance within the context of biophysical ecology (engineering methods of heat, mass and momentum

exchange applied to biological systems), such approaches will be better poised to predict where and when the impacts of climate change will most likely occur. Hendee, JC and Berkelmans, R. (2000) Expert system generated coral bleaching alerts for Myrmidon and Agincourt Reefs, Great Barrier Reef, Australia. Proceedings of the 9th International Coral Reef Symposium: Bali (Indonesia), 1099-1104. An expert system, termed the Coral Reef Early Warning System (CREWS), was employed at NOAA’s Atlantic Oceanographic and Meteorological Laboratory (AOML) in Miami, Florida, USA, to provide interpretations of combinations of near real-time meteorological and oceanographic data, thought to be conducive to coral bleaching. These data were collected via HF radio from automatic weather stations operated by the Australian Institute of Marine Science at Myrmidon and Agincourt Reefs in the central Great Barrier Reef. At Myrmidon Reef, CREWS was useful in “predicting” coral during the end of January and early February 2000. This prediction was verified in the field with observations of low level bleaching for a few species of Acropora bleaching (especially Acropora gemmifera and A. digitifera) on the reef flat at Myrmidon Reef. In February 2000, alerts were produced and sent for Agincourt Reef, 330km north of Myrmidon Reef, but field observations showed that bleaching did not occur. These results suggest that CREWS may be fine-tuned to predict bleaching in thermally sensitive species or genera at individual locations. Hendee, JC, Mueller, E, Humphrey, C and Moore, T. (2001) A data-driven expert system for producing coral bleaching alerts at Sombrero Reef in the Florida Keys. Bulletin of Marine Science 69, 673-684. A computer expert system shell was employed to provide interpretations of near real-time acquired combinations of meteorological and oceanographic parameters from a SEAKEYS (Sustained Ecological Research Related to Management of the Florida Keys Seascape) station at Sombrero Reef. When environmental conditions were conducive to coral bleaching, according to different models, alerts were automatically posted to the World-Wide Web and emailed to researchers so they could verify and study bleaching events as they might happen. The models were refined using feedback from field data on bleaching recorded after alerts from the expert system. The expert system was programmed to produce alerts when sea temperatures over 30°C occurred, or when temperatures of 30°C occurred concomitant with low winds. Alerts were produced in June 1998 when these conditions were met, but bleaching did not occur. Reconfiguration of the system, which included a point system for three models (high sea temperature only, high sea temperature plus low winds, high sea temperature plus low winds plus low tide), resulted in the transmittal of alerts which coincided with bleaching during early August 1998. Bleaching occurred after sea temperature reached an average of 31.5°C over a period of three days, with excursions over 31.8°C occurring over 15 times during those three days. High sea temperatures, low wind speeds, and a very low tide occurred coincident to the time of bleaching, but it was not possible to tell if these were factors acting synergistically.

Hendee, JC (ed.) (2004) The effects of combined sea temperature, light, and carbon dioxide on coral bleaching, settlement, and growth. First Annual Combined Effects Think Tank to Support CREW Modeling: Lee Stocking Island (Bahamas), 126 pp. Researchers engaged in studies of coral bleaching, growth, reproduction and other disciplines gathered at the Caribbean Marine Research Center (CMRC) on Lee Stocking Island, Bahamas (site of the first Coral Reef Early Warning System, or CREWS stations) January 20-24, 2003, to discuss how their research problems might gain from a supplemental CREWS approach, and how the CREWS knowledge base could gain from their work. The objectives of the meeting were to, a) explain what is known of certain coral reef problem domains, that is, as they relate to anthropogenic stress and coral reef response, b) determine which in situ monitoring instruments would help in these research problems, and, c) to begin to provide a dialog to enhance current expert system modules, or develop new ones, to facilitate interpretation of (i.e., model) those factors thought to be conducive to stimulating a particular event (e.g., coral bleaching, coral growth). The work presented at this Think Tank represents a milestone in marking the status of important topics in coral reef research today, and indeed new research directions have already begun. The present NOAA Technical Memorandum details the salient points of the invitees' presentations through summaries, relevant bibliographic citations, and links (at the bottom of the first page of each presentation summary) to their PowerPoint presentations on the World-Wide Web. The document is arranged to show the meeting agenda followed by the presentations in the order in which they were given. Based on the importance of CREWS research, we expect to conduct future Think Tank meetings to summarize current findings and identify important new coral reef research topics. Henderson, C. (2006) Paradise lost. New Scientist, 28-33. As the oceans turn acid, coral reefs will vanish along with innumerable other sea creatures. Caspar Henderson reports on an disaster in the making Heyward, AJ, Smith, LD, Rees, M, Field, SN. (2002) Enhancement of coral recruitment by in situ mass culture of coral larvae. Marine Ecology Progress Series 230, 113-118. New technologies for culturing and settling scleractinian coral larvae in the field are required to elucidate the role of recruitment in population dynamics and to provide options for reef rehabilitation. Natural multi-species aggregations of coral embryos, which frequently form slicks on the sea surface after large-scale annual spawnings, were identified as a potential resource for mass coral culture. Slicks containing billions of embryos were found at sea and several million embryos were sub-sampled and successfully cultured in simple floating ponds, moored over the reef. Coral larvae were maintained in the floating ponds until competent to settle, and then seeded onto the reef environment via hoses from the ponds to mesh enclosures temporarily fixed to the reef substratum. Reefal areas exposed to the cultured larvae exhibited a dramatic enhancement of coral larval recruitment compared to natural rates. The results demonstrate the ability to seed defined areas of reef with controlled densities of recruits. We conclude that natural spawning slicks, which have been noted on

numerous reefs throughout the world, provide opportunities for very large-scale culture of corals, which may have application in reef rehabilitation and management strategies where natural recruitment is limited. Hill, J and Wilkinson, C. (2004) Methods for ecological monitoring of coral reefs. Australian Institute of Marine Science: Townsville (Australia), 117 pp. The aim of this book is to help managers of coral reefs select appropriate ecological monitoring programs, protocols and methods for your coral reef management needs. This book was written in response to requests from coral reef managers for advice on monitoring, especially: - How monitoring can help management; - How to choose the best methods to suit your needs; and - The good and bad points and associated costs of a wide range of monitoring methods. Monitoring can be specific or general. There are different management information needs for each coral reef area, so monitoring programs must be designed to include a selection of protocols and methods to meet those needs. The protocols and methods outlined in this book represent the ones most commonly used on coral reefs around the world. Our advice is to use the standard and frequently used methods to monitor your reefs because these have been extensively tested. Using standard methods also means that you will be able to compare the status of your coral reefs with other reefs at regional and global scales. Coral reef managers around the world have similar problems and questions that monitoring can answer. For example, managers need to know if: - Coral reefs are healthy and improving; - Threats are damaging the corals or other organisms; - Fish populations are increasing in a protected area; - Management actions have been successful; - Economies of local communities are maintained or improved; - Communities understand the need for management and want to assist; - Tourism is a positive or negative benefit for the coral reef area. These questions and many others can be answered with an effective monitoring program, which will consist of a number of monitoring methods, often at a mix of scales from the whole reef to a small area. This reference book should be kept current. We invite you to recommend additional methods to be included as well as other suggested updates. Please write to us at c.wilkinson@aims.gov.au. Hill, R, Larkum, AWD, Frankart, C, Kühl, M and Ralph, PJ. (2004) Loss of functional Photosystem II reaction centres in zooxanthellae of corals exposed to bleaching conditions: Using fluorescence rise kinetics. Photosynthesis Research 82, 59-72. Mass coral bleaching is linked to elevated sea surface temperatures, 1–2 °C above average, during periods of intense light. These conditions induce the expulsion of zooxanthellae from the coral host in response to photosynthetic damage in the algal symbionts. The mechanism that triggers this release has not been clearly established and to further our knowledge of this process, fluorescence rise kinetics have been studied for the first time. Corals that were exposed to elevated temperature (33 °C) and light (280 µmol photons m-2 s-1), showed distinct changes in the fast polyphasic

induction of chlorophyll-a fluorescence, indicating biophysical changes in the photochemical processes. The fluorescence rise over the first 2000 ms was monitored in three species of corals for up to 8 h, with a PEA fluorometer and an imaging-PAM. Pocillopora damicornis showed the least impact on photosynthetic apparatus, while Acropora nobilis was the most sensitive, with Cyphastrea serailia intermediate between the other two species. A. nobilis showed a remarkable capacity for recovery from bleaching conditions. For all three species, a steady decline in the slope of the initial rise and the height of the J-transient was observed, indicating the loss of functional Photosystem II (PS II) centres under elevated temperature conditions. A significant loss of PS II centres was confirmed by a decline in photochemical quenching when exposed to bleaching stress. Non-photochemical quenching was identified as a significant mechanism for dissipating excess energy as heat under the bleaching conditions. Photophosphorylation could explain this decline in PS II activity. State transitions, a component of non-photochemical quenching, was a probable cause of the high non-photochemical quenching during bleaching and this mechanism is associated with the phosphorylation-induced dissociation of the light harvesting complexes from the PS II reaction centres. This reversible process may account for the coral recovery, particularly in A. nobilis. Hill, R, Schreiber, U, Gademann, R, Larkum, AWD, Kühl, M and Ralph, PJ. (2004) Spatial heterogeneity of photosynthesis and the effect of temperature-induced bleaching conditions in three species of corals. Marine Biology 144, 633-640. Heterogeneity in photosynthetic performance between polyp and coenosarc tissue in corals was shown using a new variable fluorescence imaging system (Imaging-PAM) with three species of coral, Acropora nobilis, Cyphastrea serailia and Pocillopora damicornis. In comparison to earlier studies with fibre-optic microprobes for fluorescence analysis, the Imaging-PAM enables greater accuracy by allowing different tissues to be better defined and by providing many more data points within a given time. Spatial variability of photosynthetic performance from the tip to the distal parts was revealed in one species of branching coral, A. nobilis. The effect of bleaching conditions (33°C vs. 27°C) was studied over a period of 8 h. Marked changes in fluorescence parameters were observed for all three species. Although a decline in ΦPSII (effective quantum yield) and Yi (the first effective quantum yield obtained from a rapid light curve) were observed, P. damicornis showed no visual signs of bleaching on the Imaging-PAM after this time. In A. nobilis and C. serailia, visual signs of bleaching over the 8 h period were accompanied by marked changes in F (light-adapted fluorescence yield), NPQ (non-photochemical quenching) and Ek (minimum saturating irradiance), as well as ΦPSII and Yi. These changes were most marked over the first 5 h. The most sensitive species was A. nobilis, which after 8 h at 33°C had reached a ΦPSII value of almost zero across its whole surface. Differential bleaching responses between polyps and coenosarc tissue were found in P. damicornis, but not in A. nobilis and C. serailia. NPQ increased with exposure time to 33°C in both the latter species, accompanied by a decreasing Ek, suggesting that the xanthophyll cycle is entrained as a mechanism for reducing the effects of the bleaching conditions.

Hill, R, Frankart, C and Ralph, PJ. (2005) Impact of bleaching conditions on the components of non-photochemical quenching in the zooxanthellae of a coral. Journal of Experimental Marine Biology and Ecology 322, 83-92. Mass coral bleaching events are a worldwide phenomenon, which generally occur during periods of elevated sea surface temperature and intense sunlight. These conditions result in a decline in photochemical efficiency of symbiotic microalgae (zooxanthellae) which ultimately leads to the expulsion of these symbionts. The physiological mechanism which triggers the release of the zooxanthellae has yet to be adequately determined. Under bleaching conditions, non-photochemical quenching (NPQ) is used to dissipate excess energy from photosystem II (PSII). NPQ was partitioned into three components, (energy dependent quenching [qE], state transition quenching [qT] and photoinhibitory quenching [qI]), based on relaxation kinetics upon addition of 3-(3,4-dichlorophenyl)-1,1-dimethylurea (DCMU) and darkening. This investigation revealed that for corals not exposed to bleaching stress, qE was the principle means of energy dissipation (~60% of the total NPQ). In corals exposed to either high-light (475 µmol photons m-2 s-1 and 25°C) or elevated temperature (225 µmol photons m-2 s-1 and 32°C) treatments, the dominant component of NPQ was qE and the relative proportions did not change during the exposure period (1–8 h). When exposed to bleaching conditions (475 µmol photons m-2 s-1 and 32°C) the contribution of the different components changed after 4 h and the total NPQ increased. At this time, the contribution of qT to the total NPQ significantly increased to equal that of qE (40%), suggesting state transitions become more important under such conditions. Throughout the exposure period in all treatments, no change in the proportion of qI was observed. Hill, R and Ralph, J. (2006) Photosystem II Heterogeneity of in hospite Zooxanthellae in Scleractinian Corals Exposed to Bleaching Conditions. Photochemistry and Photobiology 82, 1577–1585. Increased ocean temperatures are thought to be triggering mass coral bleaching events around the world. The intracellular symbiotic zooxanthellae (genus Symbiodinium) are expelled from the coral host, which is believed to be a response to photosynthetic damage within these symbionts. Several sites of impact have been proposed, and here we probe the functional heterogeneity of Photosystem II (PSII) in three coral species exposed to bleaching conditions. As length of exposure to bleaching conditions (32°C and 350 µmol photons m-2 s-1) increased, the QA

- reoxidation kinetics showed a rise in the proportion of inactive PSII centers (PSIIX), where QB was unable to accept electrons. PSIIX contributed up to 20% of the total PSII centers in Pocillopora damicornis, 35% in Acropora nobilis and 14% in Cyphastrea serailia. Changes in FV/FM and amplitude of the J step along fast induction curves were found to be highly dependent upon the proportion of PSIIX centers within the total pool of PSII reaction centers. Determination of PSII antenna size revealed that under control conditions in the three coral species up to 60% of PSII centers were lacking peripheral light-harvesting complexes (PSIIβ). In P. damicornis, the proportion of PSIIβ increased under bleaching conditions and this could be a photoprotective mechanism in response to excess light. The rapid increases in PSIIX and PSIIβ observed in these corals under bleaching conditions indicates these physiological processes are involved in the initial photochemical damage to zooxanthellae.

Hoegh-Guldberg, O. (1999) Climate change, coral bleaching and the future of the world’s coral reefs. Marine and Freshwater Research 50, 839-866. Sea temperatures in many tropical regions have increased by almost 1°C over the past 100 years, and are currently increasing at ~1.2°C per century. Coral bleaching occurs when the thermal tolerance of corals and their photosynthetic symbionts (zooxanthellae) is exceeded. Mass coral bleaching has occurred in association with episodes of elevated sea temperatures over the past 20 years and involves the loss of the zooxanthellae following chronic photoinhibition. Mass bleaching has resulted in significant losses of live coral in many parts of the world. This paper considers the biochemical, physiological and ecological perspectives of coral bleaching. It also uses the outputs of four runs from three models of global climate change which simulate changes in sea temperature and hence how the frequency and intensity of bleaching events will change over the next 100 years. The results suggest that the thermal tolerances of reef-building corals are likely to be exceeded every year within the next few decades. Events as severe as the 1998 event, the worst on record, are likely to become commonplace within 20 years. Most information suggests that the capacity for acclimation by corals has already been exceeded, and that adaptation will be too slow to avert a decline in the quality of the world.s reefs. The rapidity of the changes that are predicted indicates a major problem for tropical marine ecosystems and suggests that unrestrained warming cannot occur without the loss and degradation of coral reefs on a global scale. Hoegh-Guldberg, O, Hoegh-Guldberg, H, Stout, DK, Cesar, H and Timmermann, A. (2000) Pacific in peril: Biological, economic and social impacts of climate change on Pacific coral reefs. Greenpeace, 72 pp. Climate change may be the single most important challenge to human societies in the 21st century. Changes to the environmental factors that govern the earth’s biological systems have flow-on effects for almost every aspect of human societies. Coral reef ecosystems are especially susceptible to climate change and recent predictions have suggested that coral reefs will be seriously degraded by the changing conditions of the world’s tropical oceans. This report follows a paper published last year by one of its authors, Professor Hoegh-Guldberg. In the first two sections, it extends and updates the previous findings on coral reefs with a closer focus on Pacific reefs. It then draws upon the physical and social sciences for an answer to the following basic question: What do the impact of climate change on coral reefs and associated climate-change effects mean for the medium- and long-term future of the people of the Pacific? The answer and a consequent policy response are needed urgently. The timeline is short and everyday policy-making concerning the current election horizon or next year tends to dominate the attention of politicians. National leaders and international organisations must be persuaded to have in place development planning responses to extend the horizon and allow the necessary fundamental policy changes to occur. The socioeconomic impacts identified in the paper are so profound that they dwarf any strategic issue currently confronting a major peacetime economy. There is a need for urgent action on an international scale. The fate of these nations and the surrounding ocean is of worldwide concern. Coral reef ecosystems are particularly vulnerable to changes in sea temperature, sea level and oceanic

alkalinity. Increasing episodes of coral bleaching have been followed by larger scale mortality events, often totally removing reef-building corals from some locations. The last major bleaching event (1998, the largest in history) saw the removal of large areas of reefbuilding corals. A major decline in the condition of the corals that build coral reefs is now a distinct possibility within three to five decades. This will have major effects on marine ecosystems along the coastlines of many tropical countries. Reef-building corals are central to the health of these ecosystems. They provide much of the primary productivity and framework within which many important marine species live and the reefs they build form crucial barriers to the open ocean. Pacific societies have been identified as being particularly vulnerable to climate change in previous studies. The impact of climate change on Pacific economies is complex and requires a truly multidisciplinary focus. The present study isolates one component of the effect of climate change, the loss of coral reefs, and assembles economic and associated data to help assess the ramifications for each of 13 Pacific nations. While the task was ambitious (and further data collection will probably refine the conclusions), the trends and conclusions within the analysis are clear. The 13 nations fall into two groups, of which four Melanesian nations are relatively resilient to change and the remaining mainly Polynesian nations generally less resistant and more vulnerable. By 2020, all 13 nations, wherever situated, are likely to face increasingly bleak conditions. Most scientists agree that the current rate of warming of the world’s oceans is a consequence of increasing concentrations of greenhouse gases in the atmosphere. The results of this study reveal that the Pacific ocean is no exception; it warmed at a mean rate of 0.79ºC over the past century. A complex geographic pattern was associated with the distribution of warming rates. Notably, Tuvalu, Kiribati, Samoa, Vanuatu and New Caledonia had warming rates higher than 1ºC per century while Tonga, the Cook Islands (southern section) and Solomon Islands experienced minimal warming when considered over the last century. Mass coral bleaching events have increased in the Pacific and have been triggered (as they have in all other coral reef domains) by 1-2ºC thermal anomalies above the average summertime maximal sea temperatures. Recent mass bleaching events in the Pacific (eg, Cook Islands in 1994, Palau in 1998, Fiji in 2000) have seen large coral mortality events among previously rich coral reef communities. Some sites have lost all of their living coral cover, which has not returned after almost 10 years. The available models for projecting climate change are described in detail in the report, including hindcasting to show that these models successfully describe events over the 20th century. The analysis reveals that the level of thermal stress has increased significantly. The thermal stress necessary to produce large-scale bleaching events is only apparent from the late 1960s onwards. Projections of bleaching frequencies and intensities were prepared for the 13 Pacific nations. The projected changes to the frequency and intensity of mass coral bleaching events are a major cause for concern. Thermal anomalies required to produce severe mass bleaching and mortality events occur in the first few decades of the 21st century. These rapidly rise to much higher values by the middle to late part of the century. There is no evidence that reefs will be able to survive these levels of thermal stress. Indeed, all evidence is to the contrary, for example, mass mortality events in Palau and Okinawa in 1998. To estimate significant dates for the health of coral reefs faced with unrestrained warming, a series of criteria was established. The dates at which three severe

mortality events occur are critical since reefs that undergo mortality events where corals are largely removed from the reef communities are extremely unlikely to become repopulated within the three-to-four-year gap between consecutive events. These dates and the summary effect of the events occur between 2010 and 2060 (mean decade 2040) if a model (assuming no aerosol component) is applied for the 13 Pacific nations studied. The dates are slightly extended (by a decade) if an aerosol component is added. Great coherence between different general circulation models (GCMs) is also seen. The stress required to trigger a bleaching event becomes an annual occurrence in the waters surrounding the 13 Pacific nations between the dates of 2010 and 2070 (mean decade 2050). Put in the context of other factors such as increased human pressure, rising sea levels, more frequent cyclones and decreased alkalinity, these changes to the survivorship of reef-building corals are likely to remove corals as dominant organisms on coral reefs in the next 20 to 50 years. How these changes to coral reefs will impact on Pacific societies is the key question analysed in the rest of the study. To understand the relative vulnerability of Pacific nations to the projected changes associated with climate change, a range of physical, demographic, social, economic, export and tourism indicators have been explored for each of the 13 nations, to assess their vulnerability to change and capacity to adapt. Not surprisingly, this analysis shows that the most vulnerable countries are Tuvalu and Kiribati, tiny islands in a vast surrounding ocean. Generally, the Polynesian nations tend to be small and vulnerable, with relatively limited possibilities to adapt to changed circumstances. The four Melanesian nations are larger and more adaptable to climate change than the large number of small, lowlying nations, which typically have monochromatic economies and a narrow range for adaptation. Beyond 2020, however, every nation still functioning faces economic decline according to the scenarios developed for the report. Given the potential impact of climate change on Pacific societies based on even mild climate-change scenarios, planning for these futures is a top priority. Planning must begin now to deal with the growing vulnerability of coastal dwellings and with other impacts on societies in which the majority of the populations depend on subsistence living, based on access to healthy reef systems and associated produce. International action is clearly required to secure the funding and technical assistance required to protect and, ultimately, rehabilitate the Pacific community of nations, once described as “pearls of priceless beauty and an earthly paradise”. Hoegh-Guldberg, O, Jones, RJ, Ward, S and Loh, WK. (2002) Is coral bleaching really adaptive? Nature 415, 601-602. From an experiment in which corals are transplanted between two depths on a Panamanian coral reef, Baker infers that bleaching may sometimes help reef corals to survive environmental change. Although Baker's results hint at further mechanisms by which reef-building corals may acclimatize to changing light conditions, we do not consider that the evidence supports his inference.

Hoegh-Guldberg, O and Fine, M. (2004) Low temperatures cause coral bleaching. Coral Reefs 23, 444. Coral bleaching involves the loss of brown algal symbiont and is symptomatic of stress in reef-building corals. Mass coral bleaching (affecting thousands of square kilometers of coral reef) has been reported with growing frequency over the past two decades and is triggered by anomalously high sea temperatures. Here we report bleaching triggered by anomalously low sea temperatures in late July 2003. These observations are consistent with laboratory experiments in which anemones (Steen et al. 1987) and corals (Saxby et al. 2003) bleach in response to low water temperatures. Hoegh-Guldberg, O and Hoegh-Guldberg, H. (2004) The implications of climate change for Australia’s Great Barrier Reef: People and industries at risk. WWF Australia: Sydney (Australia), 12 pp. WWF Australia and the Queensland Tourism Industry Council commissioned a revolutionary new study on Australia’s Great Barrier Reef. This study explores possible ‘futures’ for the Reef, a complex task that involves the analysis of vast sets of information ranging from the climate sciences to the biology and economics of Australia’s coral reef industries. The analysis reveals a range of possible futures, some of which are more desirable than others from economic, social and environmental perspectives. Some of these futures are disturbing and must be avoided. This is the first study of its kind to outline how the future might unfold and how specific actions taken today will craft tomorrow’s world. Strong action to increase ecological resilience while simultaneously reducing greenhouse gas emissions today will be instrumental in securing the economic and environmental wealth of Australia tomorrow. Australia must act to influence the global agenda, and this report provides some guidance on what to do. The report seeks to answer the burning questions: What will climate change do to Australia’s Great Barrier Reef and its people, and what can we do to minimise future impacts? Hoey, AS and Bellwood, DR. (2007) Cross-shelf variation in the role of parrotfishes on the Great Barrier Reef. Coral Reefs. Herbivorous fishes are a key functional group on coral reefs. These fishes are central to the capacity of reefs to resist phase shifts and regenerate after disturbance. Despite this importance few studies have quantified the direct impact of these fishes on coral reefs. In this study the roles of parrotfishes, a ubiquitous group of herbivorous fishes, were examined on reefs in the northern Great Barrier Reef. The distribution of 24 species of parrotfish was quantified on three reefs in each of three cross-shelf regions. Functional roles (grazing, erosion, coral predation and sediment reworking) were calculated as the product of fish density, bite area or volume, bite rate, and the proportion of bites taken from various substrata. Inner-shelf reefs supported high densities but low biomass of parrotfishes, with high rates of grazing and sediment reworking. In contrast, outer-shelf reefs were characterised by low densities and high biomass of parrotfish, with high rates of erosion and coral predation. Mid-shelf reefs displayed moderate levels of all roles examined. The

majority of this variation in functional roles was attributable to just two species. Despite being rare, Bolbometopon muricatum, the largest parrotfish species, accounted for 87.5% of the erosion and 99.5% of the coral predation on outer-shelf reefs. B. muricatum displayed little evidence of selectivity of feeding, with most substrata being consumed in proportion to their availability. In contrast, the high density of Scarus rivulatus accounted for over 70% of the total grazing and sediment reworking on inner-shelf reefs. This marked variation in the roles of parrotfishes across the continental shelf suggests that each shelf system is shaped by fundamentally different processes. Hughes, TP, Baird, AH, Dinsdale, EA, Moltschaniwskyj, Pratchett, MS, Tanner, JE and Willis, BL. (1999) Patterns of recruitment and abundance of corals along the Great Barrier Reef. Nature 397, 59-63. Different physical and biological processes prevail at different scales. As a consequence, small-scale experiments or local observations provide limited insights into regional or global phenomena. One solution is to incorporate spatial scale explicitly into the experimental and sampling design of field studies, to provide a broader, landscape view of ecology. Here we examine spatial patterns in corals on the Great Barrier Reef, across spectrum of scales ranging from metres to more than 1,700 km. Our study is unusual because we explore large-scale patterns of a process (recruitment by juveniles) as well as patterns of adult abundance, revealing the relationship between the two. We show that coral-reef assemblages that are similar in terms of abundance may nonetheless show profound differences in dynamics and turnover, with major implications for their ecology, evolution and management. Hughes, TP and Connell, JH. (1999) Multiple stressors on coral reefs: A long-term perspective. Limnology and Oceanography 44, 932-940. Coral reefs are subject to a high frequency of recurrent biological and physical disturbances. The temporal and spatial scales of these are often large and difficult to study, so that most of our knowledge of disturbances on coral reefs comes from investigations conducted at one or a few sites, over short periods of time. We argue that studying single events in isolation can be misleading and that a longer term approach is necessary for understanding the responses of coral reef assemblages to multiple stressors. We present first a brief review of the impacts of physical disturbance (e.g., cyclones, hurricanes) on the community dynamics of coral reefs, with special attention to the effects of recurrent events. We then examine two unusually detailed, long-term data sets from Heron Island, Australia, and Jamaica which demonstrate some of the complexities of multiple stressors (broadly defined as natural or man-made disturbances). Both case studies illustrate that the effect of a particular disturbance often depends critically on the impact of previous perturbations. Consequently, even the same type of recurrent stressor can have different effects at different times, depending on history. Accordingly, when the added dimension of time is considered, the distinction between single and multiple stressors becomes blurred. Even a single event such as a hurricane can be viewed mechanistically as a multiple stressor, with short- and long-term impacts. We emphasize that multiple stressors often have significant effects on recruitment and regenerative processes of assemblages. These impacts are much less obvious than

catastrophic or chronic mortality, but they play a crucial role in community dynamics over longer time scales. Importantly, chronic anthropogenic impacts can impede the ability of coral assemblages to recover from natural disasters, even where there is little detectable effect on rates of adult mortality. Once a reef has been degraded, it is usually impossible to ascertain retrospectively the precise mechanisms that were involved or the relative importance of different events. A single survey will provide a snapshot of the status of coral reefs, but a longer term approach is required to understand the processes underlying changes in assemblages. Hughes, TP, Baird, AH, Dinsdale, EA, Tanner, JE, Moltschaniwskyj, NA, Pratchett, MS and Willis, BL. (2000) Supply-side ecology works both ways: The link between benthic adults, fecundity, and larval recruits. Ecology 81, 2241-2251. "Supply-side" ecology recognizes the potential role that recruitment plays in the local population dynamics of open systems. Apart from the applied fisheries literature, the converse link between adults and the production of cohorts of recruits has received much less attention. We used a hierarchical sampling design to investigate the relationships between adult abundance, fecundity, and rates of larval recruitment by acroporid corals on 33 reefs in five sectors (250-400 km apart) stretching from north to south along the length of the Great Barrier Reef, Australia. Our goal was to quantify patterns of recruitment at multiple scales, and to explore the underlying mechanisms. Specifically, we predicted that large-scale patterns of recruitment could be driven by changes in the abundance of adults and/or their fecundity, i.e., that corals exhibit a stock-recruitment relationship. The amount of recruitment by acroporids in each of two breeding seasons varied by more than 35-fold among the five sectors. Adult density varied only twofold among sectors and was not correlated with recruitment at the sector or reef scale. In contrast, fecundity levels (the proportion of colonies on each reef that contained ripe eggs) varied from 15% to 100%, depending on sector, year, and species. Spatial and temporal variation in the fecundity of each of three common Acropora species explained most of the variation (72%) in recruitment by acroporids, indicating that the production of larvae is a major determinant of levels of recruitment at large scales. Once fecundity was accounted for, none of the other variables we examined (sector, reef area, abundance of adults, or year) contributed significantly to variation in recruitment. The relationship between fecundity and recruitment was nonlinear, i.e., rates of recruitment increased disproportionately when and where the proportion of gravid colonies approached 100%. This pattern is consistent with the hypothesis that enhanced fertilization success and/or predator satiation occurs during mass-spawning events. Furthermore, it implies that small, sublethal changes in fecundity of corals could result in major reductions in recruitment. Hughes, TP and Tanner, JE. (2000) Recruitment failure, life histories, and long-term decline of Caribbean corals. Ecology 81, 1-17. Population decline, local extinction, and recovery are profoundly influenced by variation in demography and life-history traits. In open populations, changes in patterns of recruitment may also have a major influence on the size of local populations, particularly for short-lived organisms. We examine here the demographic processes underlying a slow decline of corals on Jamaican reefs,

where coral cover has decreased by fourfold over a 16-yr period. We divided the study into three approximately equal intervals (1977-1982, 1982-1987, and 1987-1993) and constructed size-based transition matrices for each of three abundant species of corals (Montastrea annularis, Agaricia agaricites, and Leptoseris cucullata) that differ substantially in life history: Montastrea is slower-growing, longer-lived, and has lower rates of recruitment than the other two species. Rates of survival, population growth ([lambda]), and recruitment declined substantially over time for all species and the stable size structures bec ame increasingly dominated by small colonies. Elasticity and life table response analysis showed that changes in the persistence of large colonies had the biggest impact on population growth in all species. Simulations indicated that the levels of larval recruitment required to maintain populations at 1977 levels increased sharply over time, even as the actual recruitment rate declined, Recruitment failure was much more important to A. agaricites and L. cucullata than to M. annularis, which could survive long periods with minimal larval input. Recovery of these populations will require an increase in both survival and recruitment. The likelihood of the latter will depend on the scale of larval dispersal, and on the impact of large-scale mortality of adults on stock-recruitment relationships. Differences in connectivity and life histories of corals will determine future patterns of recovery or further decline. Hughes, TP, Bellwood, DR and Connolly, SR. (2002) Biodiversity hotspots, centres of endemicity, and the conservation of coral reefs. Ecology Letters 5, 775-784. On land, biodiversity hotspots typically arise from concentrations of small-range endemics. For Indo-Pacific corals and reef fishes, however, centres of high species richness and centres of high endemicity are not concordant. Moreover ranges are not, on average, smaller inside the Central Indo-Pacific (CI-P) biodiversity hotspot. The disparity between richness and endemicity arises because corals and reef fishes have strongly skewed range distributions, with many species being very widespread. Consequently, the largest ranges overlap to generate peaks in species richness near the equator and the CI-P biodiversity hotspot, with only minor contributions from endemics. Furthermore, we find no relationship between the number of coral vs. fish endemics at locations throughout the Indo-Pacific, even though total richness of the two groups is strongly correlated. The spatial separation of centres of endemicity and biodiversity hotspots in these taxa calls for a two-pronged management strategy to address conservation needs. Hughes, TP, Baird, AH, Bellwood, DR, Card, M, Connolly, SR, Folke, C, Grosberg, R, Hoegh-Guldberg, O, Jackson, JB, Kleypas, J, Lough, JM, Marshall, P, Nyström, M, Palumbi, SR, Pandolfi, JM, Rosen, B and Roughgarden, J. (2003) Climate change, human impacts, and the resilience of coral reefs. Science 301, 929-933. The diversity, frequency, and scale of human impacts on coral reefs are increasing to the extent that reefs are threatened globally. Projected increases in carbon dioxide and temperature over the next 50 years exceed the conditions under which coral reefs have flourished over the past half-million years. However, reefs will change rather than disappear entirely, with some species already showing far greater tolerance to climate change and coral bleaching than others. International integration

of management strategies that support reef resilience need to be vigorously implemented, and complemented by strong policy decisions to reduce the rate of global warming. Hughes, TP, Bellwood, DR, Folke, C, Steneck, RS and Wilson, J. (2005) New paradigms for supporting the resilience of marine ecosystems. Trends in Ecology and Evolution 20, 380-386. Resource managers and scientists from disparate disciplines are rising to the challenge of understanding and moderating human impacts on marine ecosystems. Traditional barriers to communication between marine ecologists, fisheries biologists, social scientists and economists are beginning to break down, and the distinction between applied and basic research is fading. These ongoing trends arise, in part, from an increasing awareness of the profound influence of people on the functioning of all marine ecosystems, an increased focus on spatial and temporal scale, and a renewed assessment of the role of biodiversity in the sustainability of ecosystem goods and services upon which human societies depend. Here, we highlight the emergence of a complex systems approach for sustaining and repairing marine ecosystems, linking ecological resilience to governance structures, economics and society. Hughes, TP, Bellwood, DR, Folke, CS, McCook, LJ and Pandolfi, JM. (2006) No-take areas, herbivory and coral reef resilience. Trends in Ecology and Evolution. Coral reefs worldwide are under threat from various anthropogenic factors, including overfishing and pollution. A new study by Mumby et al. highlights the trophic relationships between humans, carnivorous and herbivorous fishes, and the potential role of no-take areas in maintaining vulnerable coral reef ecosystems. No-take areas, where fishing is prohibited, are vital tools for managing food webs, ecosystem function and the resilience of reefs, in a seascape setting that extends far beyond the boundaries of the reefs themselves. Hughes, TP, Rodrigues, MJ, Bellwood, DR, Ceccarelli, D, Hoegh-Guldberg, O, McCook, L, Moltschaniwskyj, N, Pratchett, MS, Steneck, RS and Willis, B. (2007) Phase Shifts, Herbivory, and the Resilience of Coral Reefs to Climate Change. Current Biology 17, 360-365. Many coral reefs worldwide have undergone phase shifts to alternate, degraded assemblages because of the combined effects of overfishing, declining water quality, and the direct and indirect impacts of climate change. Here, we experimentally manipulated the density of large herbivorous fishes to test their influence on the resilience of coral assemblages in the aftermath of regional-scale bleaching in 1998, the largest coral mortality event recorded to date. The experiment was undertaken on the Great Barrier Reef, within a no-fishing reserve where coral abundances and diversity had been sharply reduced by bleaching. In control areas, where fishes were abundant, algal abundance remained low, whereas coral cover almost doubled (to 20%) over a 3 year period, primarily because of recruitment of species that had been locally extirpated by bleaching. In contrast, exclusion of large herbivorous fishes

caused a dramatic explosion of macroalgae, which suppressed the fecundity, recruitment, and survival of corals. Consequently, management of fish stocks is a key component in preventing phase shifts andmanaging reef resilience. Importantly, local stewardship of fishing effort is a tractable goal for conservation of reefs, and this local action can also provide some insurance against larger-scale disturbances such as mass bleaching, which are impractical to manage directly. Huitric, M and McField, M. (2000) Effects of multiple disturbances on hard coral recruits in Glovers Reef Atoll’s lagoon, Belize. Beijer International Institute for Ecological Economics: Stockholm (Sweden), 20 pp. Glovers Reef Atoll, Belize, suffered two large-scale disturbances in the autumn of 1998: a 3-month bleaching event and a hurricane. The authors also participated in a large-scale field study that manually cleared macroalgae from a subset of the atoll’s lagoon patch reefs, creating an additional disturbance. Recruit populations on these patch reefs were sampled two weeks after the algal reduction, which was also during the bleaching event and prior to the hurricane (September 1998). The reefs were then resampled in September 1999. There was a significant reduction both in the density of recruits and in the proportion of bleached recruits between 1998 and 1999. Statistical analysis showed that the reduction in recruit density was not a result of the algal reduction. Benthic transects were conducted to assess coral cover change on these patch reefs. Qualitative observations suggest that the hurricane had little physical impact on these sheltered patch reefs and that declines in coral cover and recruit density were more likely attributable to bleaching. Some coral taxa appear to be especially sensitive (agaricids) or insensitive (Porites asteroides) to bleaching in both adult and recruit populations, while other genera exhibited more variation between adult and recruit responses. The variability of response also occurred on small geographic scales between different environmental habitats, illustrating the complexity of reef communities, even at geographically local scales. We conclude by discussing the possible management implications of these findings. Huppert, A and Stone, L. (1998) Chaos in the Pacific’s coral reef bleaching cycle. The American Naturalist 152, 447-459. There is no simple explanation for the unusual increase in coral reef bleaching events that have been occurring on a global scale over the last 2 decades. Recent studies focusing on this problem reveal that mass bleaching events have a strong periodic component, arising every ~3-4 yr in step with the El Nino climatic phenomenon. To explore this possibility further, we examine a simple oceanographic-ecological model designed to simulate the warm and cool phases of the Pacific Ocean cycle and gauge its effect on local coral reefs. This allows us to identify causes for localized "hot spots" in the ocean, whose high sea surface temperatures have disastrous consequences for corals. The underlying wave dynamics of the model lead to chaotic oscillations (every ~3-4 yr), which help explain the coexistence of both order and irregularity in the dynamics of mass bleaching. The model makes use of a temperature threshold mechanism-a bleaching event is triggered whenever temperature anomalies exceed a critical level. In a variable environment, the threshold mechanism is sensitive to background fluctuations, and their effects are studied by making use of a "stochastic resonance" formulation. Global climate

change and other trends in external background environmental conditions are all shown to strongly influence the distribution of mass coral bleaching events. Idjadi, JA, Lee, SC, Bruno, JF, Precht, WF, Allen-Requa, L and Edmunds, PJ. (2006) Rapid phase-shift reversal on a Jamaican coral reef. Coral Reefs 25, 209-211. Many Caribbean reefs have experienced a phase-shift in community structure, the principle features being a decline in coral cover and an increase in macroalgal biomass. However, one Jamaican reef—Dairy Bull on the north shore near Discovery Bay—is once again dominated by scleractinian corals and several key species have returned. Living coral cover at 6–8 m depth at Dairy Bull has doubled over the past 9 years and is now ~54%. The absolute cover of Acropora cervicornis was <1% in 1995, but increased to ~11% by January 2004. During this time the cover of macroalgae decreased by 90%, from 45 to 6%. We speculate that long-lived colonies of Montastraea annularis may have facilitated the recovery of this reef by providing structural refugia. Iglesias-Prieto, R, Beltrán, VH, LaJeunesse, TC, Reyes-Bonilla, H and Thomé, PE. (2004) Different algal symbionts explain the vertical distribution of dominant reef corals in the eastern Pacific. Proceedings of the Royal Society of London B Biological Sciences 271, 1757-1763. Symbiotic reef corals occupy the entire photic zone; however, most species have distinct zonation patterns within the light intensity gradient. It is hypothesized that the presence of specific symbionts adapted to different light regimes may determine the vertical distribution of particular hosts. We have tested this hypothesis by genetic and in situ physiological analyses of the algal populations occupying two dominant eastern Pacific corals, over their vertical distribution in the Gulf of California. Our findings indicate that each coral species hosts a distinct algal taxon adapted to a particular light regime. The differential use of light by specific symbiotic dinoflagellates constitutes an important axis for niche diversification and is sufficient to explain the vertical distribution patterns of these two coral species. International Ocean Commission Working Group on Coral Bleaching and Local Ecological Responses (2001) Understanding coral bleaching across four oceans. IOC/UNESCO: Heron Island, 37 pp. Coral bleaching is rapidly developing as a major problem for the health of coral reefs worldwide. Unfortunately, the level of our understanding of this phenomenon is relatively limited. Developing a good understanding of this phenomenon is a priority if management practices are to cope with the increasing levels of greenhouse-driven stress projected to occur over the next 50 years. This project uses the collective expertise of an international group to further our understanding of coral bleaching across three of the world’s major oceans. It spans molecular to ecological levels of investigation and will generate tools and techniques with a wide applicability to the problems facing both developed and developing nations.

International Ocean Commisiion Working Group on Coral Bleaching and Local Ecological Responses (2002) 1st Targeted Research Workshop: Research activities and seminar proceedings. Heron Island (Australia), 39 pp. Coral bleaching involves the damage and loss of the symbiotic dinoflagellates from corals on a scale that severely dwarfs other impacts on coral ecosystems. This fact together with the observation of the recent increase in the frequency and intensity of global bleaching events have prompted a mobilization of major scientific and management concern. . The explicit links to global climate change are driving an urgent need for information on how changes in global sea temperature, the major factor driving coral bleaching events across the globe, will affect the viability of the estimated 280,000 square kilometres of the world’s coral reef ecosystems. The potential scale of this ecological change is of major concern in both developing and developed nations. In response to this concern, UNESCO’s Intergovernmental Oceanographic Commission (IOC) established an expert Study Group focused on Coral Bleaching and Related Indicators of Coral Reef Health in 2000. The aim of this group was to integrate and develop research that will allow more reliable predictions of climate impacts and the development of better bioindicator tools for managers. At the first meeting held 9-11 April 2001 at IOC, Paris wide ranging discussions between the group and the representative of a related World Bank project led to the fusion of the group to become the IOC/UNESCO-World Bank Targeted Working Group on Coral Bleaching. The immediate goals of the reconstituted group are to identify critical gaps in our knowledge of the molecular to ecological processes involved in mass coral bleaching, and to develop specific, testable hypotheses that will be the focus of targeted investigations in four key ocean areas. The ocean areas selected are located in East Africa (Zanzibar), the Philippines (Bolinao), Eastern Australia (southern GBR) and Mexico (Puerto Morelos). In addition to targeted research activities, the work plan will involve local scientists and students in a series of activities that will range from training workshops to collaborative experiments. The project development is partially funded by a World Bank Block B grant, in addition to funding from host institutions like the University of Queensland and is designed to explore how a full work plan can be implemented to pursue a complex set of questions over 5 years. The outputs will include special research volumes, books and research reports as well as training workshops and postgraduate programs in the targeted research areas. An emphasis will be placed on technology and training exchange. The 2002 workshop on Heron Island was the first step in testing the concept of targeted research within one of the four ocean areas. It was resounding success and involved 32 scientists and 18 postgraduate students. The hypotheses tested ranged through a series that was developed during the April 2001 discussions in Paris. The coincidence of a major bleaching event across the Great Barrier Reef during the workshop led to some unusual opportunities for the targeted working group to pursue questions associated with a “natural” bleaching event. Among the highlights of this successful workshop were a major audit of symbiotic dinoflagellate strains, the discovery of new coral diseases for the GBR region, new insights into the role of cell suicide and apoptosis in bleaching and the important role of clonal variability in coral stress tolerance. The abstracts within this volume are testament to the idea that encouraging the collaboration of otherwise diverse groups is a successful stimulus to getting the research done that is so vitally needed to fill the critical knowledge gaps that currently face reef scientists, managers and users.

International Ocean Commission Working Group on Coral Bleaching and Local Ecological Responses (2006). http://www.ioc.unesco.org/coralbleaching/ Coral bleaching is rapidly developing as a major problem for the health of coral reefs worldwide. Unfortunately, the level of understanding of this phenomenon is limited. Developing a good understanding of the phenomenon is important if management practices are to be effective in minimising the detrimental impacts of coral bleaching as projected to occur over the next 50 years. The IOC/WB Working Group on Coral Bleaching and Local Ecological Responses was initiated in September 2000 with the goal to integrate, synthesize and develop global research on coral bleaching and related ecological impacts of climate change on coral ecosystems, and further new research findings into development of tools and techniques for improved observations, predictions and management interventions at national and global scales. Jimémenez, CE and Cortés, J. (2003) Coral cover change associated to El Niño, Eastern Pacific, Costa Rica, 1992-2001. Marine Ecology 24, 179-192. Changes in live and dead coral cover were documented at three localities off the Costa Rican central Pacific coast first in 1992 during the aftermath of the 1991–1992 El Niño; again in the period between 1994 and 1995, and last in January 2001. Recovery of coral communities after the 1991–1992 El Niño was expressed by a significant increase (~40 %) in 1994 of live coral cover at one locality (Manuel Antonio). A subsequent decrease (~50 %) in response to the very strong 1997–1998 episode was recorded at Manuel Antonio and Ballena, mainly due to partial tissue mortality of branching (Pocillopora spp.) and massive (Porites lobata) corals. Mortality of entire colonies associated to that event was scarce and confined to branching and nodular (Psammocora stellata) corals. This species was not found at one locality (Cambutal) in the 2001 survey and it is presumed locally extinct. The recovery of this coral and others will depend on recruits from surviving colonies in deeper waters and other coral communities in the vicinity. Within sites at a given locality, contrasting results in live coral cover variability were found. This is partially due to distinct coral assemblages, coral growth, physical exposure to tidal regime, and, related to the latter, variable duration and intensity of the warming event. In general, predominant meteorological conditions at the studied area are conducive to solar radiation (UV) stress during El Niño years and are related to changes in the atmosphere-ocean interactions in response to the warming events. Jokiel, PL, Lesser, MP and Ondrusek, ME. (1997) UV-absorbing compounds in the coral Pocillopora damicornis: Interactive effects of UV radiation, photosynthetically active radiation, and water flow. Limnology and Oceanography 42, 1468-1473. A direct relationship exists between irradiance of solar ultraviolet (UV) radiation and concentration of UV-absorbing compounds, know as mycosporine-like amino acids (MAAs), in the Hawaiian reef coral Pocillopora damicornis. However, MAA concentration is also influenced by flow regime and the irradiance of photosynthetically active radiation (PAR). High irradiances of UV radiation in reef environments are correlated with high PAR and high water velocity, because all three parameters diminish exponentially with increased depth. This correlation can also be

found along horizontal gradients on reefs. The clearest water is typically found on outer reefs growing in oceanic water. These ocean reefs typically experience high water velocities due to ocean swell when compared to the more turbid lagoon reefs exposed only to small wind-driven waves. Flow-modulated photosynthetic rate seems to be a major factor affecting the observed changes in MAA concentration when P. damicornis is grown under different flow regimes and identical irradiances of UV and PAR radiation. High PAR and high water velocity significantly enhance the effect of increased UV radiation on MAA concentration. Jokiel, PL and Brown, EK. (2004) Global warming, regional trends and inshore environmental conditions influence coral bleaching in Hawaii. Global Change Biology 10, 1627-1641. Hawaiian waters show a trend of increasing temperature over the past several decades that are consistent with observations in other coral reef areas of the world. The first documented large-scale coral bleaching occurred in the Hawaii region during late summer of 1996, with a second in 2002. The bleaching events in Hawaii were triggered by a prolonged regional positive oceanic sea surface temperature (SST) anomaly greater than 1°C that developed offshore during the time of annual summer temperature maximum. High solar energy input and low winds further elevated inshore water temperature by 1–2°C in reef areas with restricted water circulation (bays, reef flats and lagoons) and in areas where mesoscale eddies often retain water masses close to shore for prolonged periods of time. Data and observations taken during these events illustrate problems in predicting the phenomena of large-scale bleaching. Forecasts and hind-casts of these events are based largely on offshore oceanic SST records, which are only a first approximation of inshore reef conditions. The observed oceanic warming trend is the ultimate cause of the increase in the frequency and severity of bleaching events. However, coral reefs occur in shallow inshore areas where conditions are influenced by winds, orographic cloud cover, complex bathymetry, waves and inshore currents. These factors alter local temperature, irradiance, water motion and other physical and biological variables known to influence bleaching.

Jompa, J and McCook, L. (1998) Seaweeds save the reef?! Australian Institute of Marine Science: Townsville (Australia). Widespread bleaching of reef corals has recently been reported from inshore reefs in the Townsville region, apparently as a result of exceptional weather conditions and flooding during January. The likely causes of this bleaching include low salinity, high temperature, and high UV light intensity. We here report a surprising increase in coral bleaching in plots from which the normally abundant canopy of seaweeds had been experimentally removed.

Karako-Lampert, S, Katcoff, DJ, Achituv, Y, Dubinsky, Z and Stambler, N. (2004) Do clades of symbiotic dinoflagellates in scleractinian corals of the Gulf of Eilat (Red Sea) differ from those of other coral reefs? Journal of Experimental Marine Biology and Ecology 311, 301-314. The symbiotic association between corals and zooxanthellae has been a major contributing factor in the success of reef-building corals. Most of these endocellular microalgal symbionts belong to the dinoflagellate genus Symbiodinium. However, considerable genetic diversity was revealed within this taxon, as is evident in the several clades of Symbiodinium found in association with hermatypic corals all over the world. The coral reefs of Eilat (Aqaba), where winter temperature minima of 21°C are close to threshold values that prevent reef development, are among the northernmost reefs in the world. Furthermore, due to the circulation patterns of the Gulf, the extremely high evaporation, and lack of any riverine inputs, the Gulf’s waters are highly saline (40.5%). In spite of the extreme location, a high diversity of coral species has been reported in this area. In this study, using PCR, we specifically amplified zooxanthellae 18S ribosomal DNA from symbionts of 11 coral species, and analyzed it with respect to RFLP and DNA sequence. Of the several clades described from the same coral hosts in other localities, only A and C were found in the present study. Symbiodinium populations in the host examined from Eilat were different relative to other parts of the world. This distribution is discussed in relation to reproduction strategy: broadcasting versus brooding. Based on our results, we suggest that clade A is transferred through a closed system. As mass bleaching in the Gulf has never been observed, we suggest that the adaptive mechanisms presumably favoring clade diversity were not yet significant in our relatively cool area. Karako-Lampert, S, Hershkovits, G, Stambler, N, Simon-Blecher, N, Achituv, Y, Dubinsky, Z and Katcoff, DJ. (2006) Differential Gene Expression in Symbiodinium microadriaticum Clade B Following Stress. Marine Biotechnology 8, 268–274. Coral bleaching is caused by the loss of symbiont zooxanthellae and/or decrease in their pigments. Since the algal symbionts provide the energy basis for corals and whole reefs, their loss or impairment of function leads to widespread mortality. This phenomenon has been documented numerous times in recent years, and has extensively damaged coral reefs all over the world. Temperature has been found to be the major cause of bleaching, and rising sea temperatures have increased the frequency of these catastrophic episodes. To characterize the response of zooxanthellae to temperature stress at the molecular level, we used the mRNA differential display technique to monitor changes in the abundance of specific mRNA species in the cell under different temperature conditions. Axenically grown zooxanthellae were exposed to a range of temperatures (21.7, 17, 26°C) before extraction of their mRNA. Of numerous differentially expressed sequences, seven mRNA species were amplified by the polymerase chain reaction (PCR) and sequenced. One of those sequences was positively identified as encoding a multifunction cell surface aminopeptidase, dipeptidyl peptidase IV, which is active in cell matrix adhesion. Our work illustrates the power of the differential display technique as a useful tool to study the response of zooxanthellae to stressors.

Kelleher, G. (1999) Guidelines for Marine Protected Areas. IUCN: Gland (Switzerland), 107 pp. Marine Protected Areas (MPAs) are essential to conserve the biodiversity of the oceans and to maintain productivity, especially of fish stocks. Yet at present there are too few MPAs and not many of them are effectively managed. These guidelines set out the various steps a country should take to establish an effective network of MPAs. IUCN has defined an MPA as “any area of intertidal or subtidal terrain, together with its overlying water and associated flora, fauna, historical and cultural features, which has been reserved by law or other effective means to protect part or all of the enclosed environment”. There are two ways of establishing MPA systems: either as many relatively small sites, each strictly protected, or as a few large multiple-use areas which contain strictly protected areas within them. To conserve biodiversity, both approaches should occur within an effective programme of ecosystem management covering the marine ecosystem and the land areas that affect it. From the accumulated technical experience in this field, there is one general lesson which can be drawn. A crucial attribute of an MPA manager is integrity. Some managers have made the mistake of believing that they can fool some of the people some, or even all, of the time. The result is a breakdown in trust. The manager may appear to win a series of battles but in fact the eventual outcome is failure. Another key lesson is that time spent in preparation is an essential investment that will be repaid many times over. Proponents of MPAs have to show demonstrable benefits for stakeholders, and this takes time and diplomacy. Box 1 lists other lessons from experience in establishing and managing MPAs in various situations around the world. The Guidelines set out the following steps, each being the subject of a separate chapter: 1. Placing MPAs in their wider context. The high degree of linkage between land and adjoining sea, and the inter-connectivity of the oceans, require that MPAs be integrated into management regimes that deal with all human activities that affect marine life. Thus MPAs should be integrated with other policies for land use and use of the sea. It is also desirable for countries to make use of international agreements, notably UNCLOS and CBD. More international support is needed for MPAs and more attempts should be made to establish MPAs on the High Seas. 2. Developing the legal framework. In most countries, a key step will be to establish the legislation needed. This may either be enabling legislation, which allows the administration or communities to establish individual MPAs, or specific legislation establishing an MPA, usually as a large multiple-use area. The various requirements for the legislation are outlined, though the needs and context will differ from one country to another. 3. Working with relevant sectors. Many sectors of human activity affect the coast and the sea, and it is vital for those planning an MPA to work with these sectors from the earliest opportunity. Tourism often has most to gain from an MPA and can generate the greatest economic activity from it. Fisheries is the other key sector, and one with which it is most important to cooperate. Other relevant sectors include aquaculture, coastal development, agriculture, forestry, industry, defence and science.

4. Making partnerships with communities and other stakeholders. MPA management should understand the local communities that will be affected by the MPA and identify potential partners. It must listen to the many interests and seek ways to involve them as participants in resource management. It is recommended to build management partnerships using the collaborative management model, which is outlined in greater detail in Annex 1. 5. Selecting the sites for MPAs. Choosing the location and extent of MPAs involves a different emphasis to that of terrestrial protected areas. In many parts of the world, local people depend greatly on the services and resources provided by natural terrestrial areas. However, the dependence on marine areas tends to be even greater. Some forms of fishing can occur in large areas without threatening the conservation objectives of the MPA because they do not involve habitat modification. This makes it feasible to balance conservation and the needs of local people. Weight needs to be given to events outside the MPA that might affect it, such as pollution. Following these principles, the guidelines propose a rigorous set of criteria for site selection that have been applied in many countries over the past few years. 6. Planning and managing the MPA. Management should be responsive and adaptive, working with local interests in a way that builds support for the conservation objectives. To achieve this, managers should adopt a systems approach, use interdisciplinary teams and follow a clear sequence of decision-making. Most MPA management is about managing human activities, so this must be at the heart of the approach. Suggested contents for a management plan are provided in Annex 2. 7. Zoning, in which various areas are allocated for various uses. This is usually the best way of ensuring strict protection of a core zone as part of a larger, multiple-use area. The stages involved in preparing a zoning plan are outlined in Annex 3. 8. Planning for financial sustainability. Lack of funds is a critical problem for many MPAs. Managers therefore need the freedom to raise funds in as many ways as possible, such as user fees, donations and environment funds, and to retain those funds for management of the MPA. External donors are advised to extend the aid period for protected area projects, so as to help achieve financial sustainability. 9. Ensuring research, monitoring, evaluation and review. Research and monitoring should be firmly orientated to solving management issues. Guidance is given on the planning and development of a monitoring and research programme, with its different emphases in the planning and the implementation phase of the MPA. Most important of all is to use the results of research and monitoring to evaluate and if necessary reorient management. Kelmo, F, Attrill, MJ and Jones, MB. (2003) Effects of the 1997-1998 El Niño on the cnidarian community of a high turbidity coral reef system (northern Bahia, Brazil). Coral Reefs 22, 541-550. We are reporting on the results of a 4-year study that allowed for the analysis of the effects of the 1997–1998 El Niño Southern Oscillation (ENSO) event on the cnidarian community of the naturally highturbidity reefs of northern Bahia, Brazil. Cnidarian densities were significantly different between pre- and post-ENSO years, with significant consequent changes in community composition. BIOENV analysis identified variations in turbidity, mean temperature variance, and cloud cover as the factors best explaining changes in the community. We suggest that the 1997–1998 ENSO event, the most intense on record, had a differential effect on the cnidarian community. Highest mortalities were suffered by minute hydroid colonies, partial

mortality by octocorals and actiniarians, while large hydroid colonies and scleractinians were the least affected. Such an ability to survive under the stressful environmental regime imposed by this ENSO event is attributed to the presence of morphological pre-adaptations (e.g., size of colony polyps, abundance of certain nematocyst types) and the inherent phenotypic plasticity that results from long-term exposure to naturally stressful conditions. Kleypas, JA, Buddemeier, RW, Archer, D, Gattuso, J-P, Langdon, C, Opdyke, BN. (1999) Geochemical consequences of increased atmospheric carbon dioxide on coral reefs. Science 284, 118-120. A coral reef represents the net accumulation of calcium carbonate (CaCO3) produced by corals and other calcifying organisms. If calcification declines, then reef-building capacity also declines. Coral reef calcification depends on the saturation state of the carbonate mineral aragonite of surface waters. By the middle of the next century, an increased concentration of carbon dioxide will decrease the aragonite saturation state in the tropics by 30 percent and biogenic aragonite precipitation by 14 to 30 percent. Coral reefs are particularly threatened, because reef-building organisms secrete metastable forms of CaCO3, but the biogeochemical consequences on other calcifying marine ecosystems may be equally severe. Kleypas, JA, Feely, RA, Fabry, VJ, Langdon, C, Sabine, CL and Robbins, LL. (2006) Impacts of ocean acidification on coral reefs and other marine calcifiers: A guide for future research. Report of a workshop sponsored by the National Science Foundation, the National Oceanic and Atmospheric Administration and the US Geological Survey: St. Petersburg (USA), 88 pp. Research findings of the past decade have led to mounting concern that rising atmospheric carbon dioxide (CO2) concentrations will cause changes in the ocean’s carbonate chemistry system, and that those changes will affect some of the most fundamental biological and geochemical processes of the sea. Thanks to the efforts of large-scale physical and biogeochemical ocean programs such as WOCE, JGOFS, and OACES, ocean-wide changes in the carbonate system are now well documented. Since 1980 ocean uptake of the excess CO2 released by anthropogenic activities is significant; about a third has been stored in the oceans. The rate of atmospheric CO2 increase, however, far exceeds the rate at which natural feedbacks can restore the system to normal conditions. Oceanic uptake of CO2 drives the carbonate system to lower pH and lower saturation states of the carbonate minerals calcite, aragonite, and high-magnesium calcite, the materials used to form supporting skeletal structures in many major groups of marine organisms. A variety of evidence indicates that calcification rates will decrease, and carbonate dissolution rates increase, as CaCO3 saturation state decreases. This evidence comes from principles of thermodynamics, the geologic record, and the evolutionary pathways of CaCO3 secreting organisms. Further evidence, from controlled experiments of biocalcification under increased CO2 conditions, confirms that calcification rates of many organisms decrease with decreasing CaCO3 saturation state. Extrapolation of these results to the real world suggests that calcification rates will decrease up to 60% within the 21st century. We know that such extrapolations are oversimplified and do not fully consider other environmental and biological effects (e.g., rising water temperature, biological adaptation); nor do they address effects on

organism fitness, community structure, and ecosystem functioning. Any of these factors could increase or decrease the laboratory-based estimates, but it is certain that net production of CaCO3 will decrease in the future. The St. Petersburg Workshop, sponsored by NSF, NOAA, and the USGS, and held at the USGS Center for Coastal and Watershed Studies on 18–20 April 2005, was designed to take the next step toward understanding the response of marine calcification to increasing atmospheric CO2 concentration. The aims of the workshop were to summarize existing knowledge on the topic, reach a consensus on what the most pressing scientific issues are, and identify future research strategies for addressing these issues. Although workshop participants were drawn from a wide range of scientific disciplines, there was a clear convergence on the major scientific issues that should be pursued over the next 5–10 years. These include: • Determine the calcification response to elevated CO2 in benthic calcifiers such as corals (including cold-water corals), coralline algae, foraminifera, molluscs, and echinoderms; and in planktonic calcifiers such as coccolithophores, foraminifera, and shelled pteropods; • Discriminate the various mechanisms of calcification within calcifying groups, through physiological experiments, to better understand the cross-taxa range of responses to changing seawater chemistry; • Determine the interactive effects of multiple variables that affect calcification and dissolution in organisms (saturation state, light, temperature, nutrients) through continued experimental studies on an expanded suite of calcifying groups; • Establish clear links between laboratory experiments and the natural environment, by combining laboratory experiments with field studies; • Characterize the diurnal and seasonal cycles of the carbonate system on coral reefs, including commitment to long-term monitoring of the system response to continued increases in CO2; • In concert with above, monitor in situ calcification and dissolution in planktonic and benthic organisms, with better characterization of the key environmental controls on calcification; • Incorporate ecological questions into observations and experiments; e.g., How does a change in calcification rate affect the ecology and survivorship of an organism? How will ecosystem functions differ between communities with and without calcifying species? • Improve the accounting of coral reef and open ocean carbonate budgets through combined measurements of seawater chemistry, CaCO3 production, dissolution and accumulation, and, in near-shore environments, bioerosion and off-shelf export of CaCO3; • Quantify and parameterize the mechanisms that contribute to the carbonate system, through biogeochemical and ecological modeling, and apply such modeling to guide future sampling and experimental efforts; • Develop protocols for the various methodologies used in seawater chemistry and calcification measurements. Some of these research objectives require technological development, but others can be addressed immediately. While much work remains toward answering the fundamental question: “How will marine calcification rates respond to increasing atmospheric CO2 concentrations,” we need to begin investigations that look forward to answering the question: “What are the consequences of reduced calcification in both planktonic and benthic calcifying communities and ecosystems?” We should not wait until we answer the former question before tackling the latter.

This report is intended as a guide to program managers and researchers toward designing research projects that address these important questions. It is written with the detail and references needed to serve as a resource for researchers, including graduate students, who wish to tackle projects within the sometimes confusing topic of marine carbonate chemistry and calcification. Knowlton, N. (2001) The future of coral reefs. Proceedings of the National Academy of Sciences of the United States of America 98, 5419-5425. Coral reefs, with their millions of species, have changed profoundly because of the effects of people, and will continue to do so for the foreseeable future. Reefs are subject to many of the same processes that affect other human-dominated ecosystems, but some special features merit emphasis: (i) Many dominant reef

builders spawn eggs and sperm into the water column, where fertilization occurs. They are thus particularly vulnerable to Allee effects, including potential extinction associated with chronic reproductive failure. (ii) The corals likely to be most resistant to the effects of habitat degradation are small, short-lived "weedy" corals that have limited dispersal capabilities at the larval stage. Habitat degradation, together with habitat fragmentation, will therefore lead to the establishment of genetically isolated clusters of inbreeding corals. (iii) Increases in average sea temperatures by as little as 1°C, a likely result of global climate change, can cause coral "bleaching" (the breakdown of coral-algal symbiosis), changes in symbiont communities, and coral death. (iv) The activities of people near reefs increase both fishing pressure and nutrient inputs. In general, these processes favor more rapidly growing competitors, often fleshy seaweeds, and may also result in explosions of predator populations. (v) Combinations of stress appear to be associated with threshold responses and ecological surprises, including devastating pathogen outbreaks. (vi) The fossil record

suggests that corals as a group are more likely to suffer extinctions than some of the groups that associate with them, whose habitat requirements may be less stringent. Kokita, T, Nakazono, A. (2001) Rapid response of an obligately corallivorous filefish Oxymonacanthus longirostris (Monacanthidae) to a mass coral bleaching event. Coral Reefs 20, 155-158. Abstract not available. Kramer, PA and Kramer, PR. (2002) Ecoregional conservation planning for the Mesoamerican Caribbean Reef. WWF: Washington DC, 140 pp. The Mesoamerican Caribbean Ecoregion (MACR) extends over 1,000 km along the coastlines of Yucatán Mexico, Belize, Guatemala, and Honduras, and supports the second longest barrier reef in the world, a diverse array of fauna and flora, and numerous rich nursery/feeding grounds. To help guide conservation efforts toward preserving the integrity of this unique ecoregion, WWF initiated Ecoregional Conservation Planning (EC) by supporting the development of a MACR Biodiversity Database and sponsoring an Ecoregional Conservation Planning Workshop in Cancun in April 2000. A preliminary goal of the EC process is to identify a network of conservation priority sites that will represent the region’s biodiversity, while contributing to the maintenance and resilience of ecological processes. This report

aims to provide a greater understanding of the region’s biodiversity and present the results and recommendations of the EC workshop. Evaluating MACR Biodiversity To understand the MACR’s biodiversity, distinctive indicators such as key species, representative habitats, and key ecological processes were targeted for further examination. Using a Geographic Information System (GIS) as a platform, a MACR Biodiversity Database containing such information as land cover, marine benthic habitats, currents watersheds, and the distribution and status of coral reefs, focal species (manatees, sea turtles, birds) and commercially significant species like lobster, grouper, and conch was created. To promote the evaluation of large-scale features (100-1,000 kms), the ecoregion was delineated into distinctive subregions according to similar biodiversity features such as oceanographic patterns (e.g., currents, upwelling), characteristic habitat types (e.g., coral reefs, seagrass, mangroves), bathymetry, watersheds and terrestrial influences. Recognizing the vital role humans play in the maintenance of biodiversity, socioeconomic data such as human uses, threats to biodiversity, and management efforts was integrated. This information was compiled into a series of thematic 3’ x 3’ maps and used as visual conservation planning tools at the EC workshop in Cancun, Mexico (see Appendix A). Ecoregional Conservation Planning Workshop Using a priority-setting framework, the EC workshop provided more than 60 local and international experts the opportunity to evaluate biodiversity properties of the MACR ecoregion, select those areas of highest conservation priority, and recommend actions to ensure their long-term preservation. The key in adapting this process was recognizing the open and transboundary nature of this large and interconnected tropical marine ecosystem by focusing on the importance of connectivity, larval distribution, currents, habitat linkages, and upwelling. Identifying Priority Areas for Taxa/Guilds The first priority-setting analysis focused on selecting priority areas important for the conservation of significant taxa or guilds, including corals, fishes, focal species, and plants. Conserving areas important to these groups may contribute to the protection and maintenance of ecosystem processes and functions throughout the ecoregion. Taxa/Guild Priority Areas were ranked based on criteria such as uniqueness, endemism, trophic importance, representation, ecological phenomena (e.g., spawning aggregations), economic importance, vulnerability, and need for inventories in their ranking analysis. Given their predominance and importance in the MACR, ranking criteria specific to coral reefs, including habitat diversity, size, condition, proximity to similar habitats, nursery areas or larval transport, environmental gradients, well-developed coral framework, regenerative capacity, representation, “swath” potential, uniqueness, and dominance of “old growth” corals, were developed and evaluated. From these analyses, 137 areas were selected including 26 Coral Priority Areas, 53 Fish Priority Areas, 37 Focal Species Priority Areas, and 21 Plant Priority Areas. Taxa/Guild Priority Areas included important coral reefs such as the longest barrier reef in the western Atlantic (Belize) and four of the best-developed coral atolls (Banco Chinchorro, Lighthouse, Turneffe Islands, and Glovers Reef Atoll) in the Caribbean; largest snapper-grouper spawning/aggregation (2,000-4,000 individuals) (Gladden Spit); sandy beaches with some of the highest densities of sea turtle nests (e.g., Cozumel, Punta de Manabique); large manatee populations (e.g., Belize River, Northern and Southern Lagoons) and important manatee calving areas (e.g., Chetumal and Corozal Bays); and numerous large nursery grounds for key

fishery species like lobster (e.g., Bahía de la Ascención, and Espíritu Santo), conch (e.g., Banco Chinchorro), and shrimp (e.g., Graciosa Bay-Amatique Bay). An overview of the Taxa/Guild Priority Areas is provided in Appendix B. Selecting Biodiversity Priority Areas for the MACR The Taxa/Guild Priority Areas were then integrated and served as the basis for identifying Ecoregional Biodiversity Priority Areas. Experts conducted an ecoregional Biodiversity Analysis to select and rank priority areas into four categories—Highest Priority, High Priority, Priority, and Areas Requiring Additional Assessments (or Unknown)—using several criteria such as species richness, trophic linkages, habitat connectivity, habitat complexity, habitat representation, and ecological and evolutionary phenomena. Twenty-six Biodiversity Priority areas were selected: 9 Highest, 7 High, 6 Priority, and 4 Unknown (see table below). Detailed descriptions are found in Appendix C. Threats to MACR Biodiversity To evaluate threats to MACR biodiversity, a socioeconomic working group identified and ranked the primary threats to the ecoregion’s biodiversity as high, medium, or low based on four of the greatest threats: coastal habitat degradation or conversion, declining water quality, declining or depleted fisheries, and increased stresses due to oceanographic and climato-meteorological phenomena. Areas at greatest risk included Cancun (development), Chetumal Bay (pollution), Belize City (shipping, development), La Ceiba and Trujillo (agriculture, oil-port activities), and Bay Islands (development, overfishing). Identifying Persistence Likelihood and Future Threats A Persistence Analysis of the Biodiversity Areas was then conducted based on the degree to which an area would likely retain its present status should the current level of human pressure remain unchanged. Several landscape and integrity criteria were ranked, including intactness of trophic structures, linkages to adjacent ecosystems, habitat quality, structure and complexity, status of key species and susceptibility to large-scale disturbances. Of the 26 Biodiversity Priority areas, 11 were considered to have a high persistence value, 11 were ranked moderate, 2 were ranked low, and 2 were ranked as unknown. No Priority areas were ranked as having the highest likelihood of persistence. The Biodiversity Priority Areas with the greatest likelihood of persistence included NE Yucatán (QR2), Sian Ka’an (SK3), Banco Chinchorro (SK1), Lighthouse Reef (BB2), Turneffe Islands (BB1), Gladden Spit (BB6), Belize City Complex (BB4), Ría Lagartos (QR3), Central Barrier Reef (BB7), and Río Platano (HG2). The final analysis was a Future Threats Analysis where experts evaluated the likelihood that threats would result in significant changes in the next 15 years, based on three broad categories of threats: 1) landbased (e.g., pollution associated with runoff); 2) marine-aquatic (e.g., loss or disruption of nursery or spawning grounds); and 3) marine biota (e.g., shifts in community structure, unsustainable fishing). Fourteen of the Biodiversity Priority Areas were ranked as having high levels of threats, nine as moderate, only one as low, and two as unknown. Those at highest risk include: NE Yucatán (QR2), S. Yucatán (SK4), Glovers Reef Atoll (BB3), Gladden Spit (BB6), Bay Islands (HG3), Cozumel (QR1), Central Yucatán (SK2), Chetumal/Corozal (SK5), Belize City Complex (BB4), Sapodilla Cayes (BB8), Port Honduras (BB9), Gulf of Honduras (HG5), Cancun Corridor (QR5), Tulum Corridor (QR6), and Tela-Manabique Coast (HG4). Seascape Considerations The experts discussed important seascape features to consider for developing a long-term conservation plan for the MACR. They identified specific conservation

strategies and actions needed to achieve and measure conservation success over the next one to two years, five to ten years and 25-50 years. Workshop experts also identified important data needs such as status and condition of key habitats, organisms, and fisheries; water quality and sources of contamination; sustainable development, and effects of global climate change. Subsequent to the workshop, an area analysis was conducted to evaluate the range of representative habitat types within the system of Biodiversity Priority Areas. The results of the workshop are seen as an initial step to help focus conservation efforts by identifying those areas of highest priority as well as those at greatest risk. With better insight into the important areas and ecological processes of the MACR, WWF will continue to work with partners toward our common goal of preserving the region’s unique biodiversity and ensuring its continued contribution to the ecological health of the region and the livelihood of present and future generations. The relatively low state of development and overall good environmental quality of most of the ecoregion provide a unique conservation opportunity to prevent the kinds of ecological damages that have occurred in similar habitats in other parts of the world. Kuffner, IB. (2005) Temporal Variation in Photosynthetic Pigments and UV-Absorbing Compounds in Shallow Populations of Two Hawaiian Reef Corals. Pacific Science 59, 561-580. As we seek to understand the physiological mechanisms of coral bleaching, it is important to understand the background temporal variation in photosynthetic pigments and photoprotective compounds that corals exhibit. In this study, reef flat populations of two hermatypic coral species, Montipora capitata (Dana, 1846) and Porites compressa Dana, 1846, were sampled monthly in Kāne‘ohe Bay, Hawai‘i, from January 1998 to March 1999. Surface ultraviolet radiation (UVR) was measured continually during this time period at the same location. High-performance liquid chromatography (HPLC) analysis of photosynthetic pigments and mycosporine-like amino acids (MAAs) revealed temporal changes in concentrations and proportions of these compounds in tissues of both species of coral. Chlorophyll a (chl a), chlorophyll c2 (chl c2), peridinin, and diadinoxanthin concentrations changed on a skeletal weight (M. capitata) or surface area (P. compressa) basis, significantly correlating with seasonal changes in solar input (number of days from the winter solstice). In P. compressa, diadinoxanthin increased in proportion to the total pigment pool during summer months, suggesting an up-regulation of a xanthophyll cycle. In M. capitata, the ratio of chl a: chl c2 decreased during winter months, suggesting photoacclimation to lower light levels. It is surprising that there was not a clear seasonal pattern in total MAA concentration for either species, with the exception of shinorine in P. compressa. The relative stability of MAA concentrations over the course of the year despite a pronounced seasonal trend in UVR suggests either that MAAs are not performing a photoprotective role in these species or that concentrations are kept at a threshold level in the presence of a dynamic light environment.

Kuffner, IB, Walters, LJ, Becerro, MA, Paul, VJ, Ritson-Williams, R and Beach, KS. (2006) Inhibition of coral recruitment by macroalgae and cyanobacteria. Marine Ecology Progress Series 323, 107-117. Coral recruitment is a key process in the maintenance and recovery of coral reef ecosystems. While intense competition between coral and algae is often assumed on reefs that have undergone phase shifts from coral to algal dominance, data examining the competitive interactions involved, particularly during the larval and immediate post-settlement stage, are scarce. Using a series of field and outdoor seawater table experiments, we tested the hypothesis that common species of macroalgae and cyanobacteria inhibit coral recruitment. We examined the effects of Lyngbya spp., Dictyota spp., Lobophora variegata (J. V. Lamouroux) Womersley, and Chondrophycus poiteaui (J. V. Lamouroux) Nam (formerly Laurencia poiteaui) on the recruitment success of Porites astreoides larvae. All species but C. poiteaui caused either recruitment inhibition or avoidance behavior in P. astreoides larvae, while L. confervoides and D. menstrualis significantly increased mortality rates of P. astreoides recruits. We also tested the effect of some of these macrophytes on larvae of the gorgonian octocoral Briareum asbestinum. Exposure to Lyngbya majuscula reduced survival and recruitment in the octocoral larvae. Our results provide evidence that algae and cyanobacteria use tactics beyond space occupation to inhibit coral recruitment. On reefs experiencing phase shifts or temporary algal blooms, the restocking of adult coral populations may be slowed due to recruitment inhibition, thereby perpetuating reduced coral cover and limiting coral community recovery. Kuguru, B, Winters, G, Beer, S, Santos, SR and Chadwick, NE. (2007) Adaptation strategies of the corallimorpharian Rhodactis rhodostoma to irradiance and temperature. Marine Biology 151, 1287-1298. Corallimorpharians may dominate some habitats on coral reefs and compete with stony corals for access to light, yet little is known concerning their photosynthetic traits. At Eilat in the northern Red Sea, we observed that the abundance of individuals of the corallimorpharian Rhodactis rhodostoma decreased significantly with depth on the reef slope. Field and laboratory experiments revealed that they employ several mechanisms of photoadaptation to high irradiance on the shallow reef Xat. Their endosymbiotic microalgae (zooxanthellae) varied significantly in both abundance and chlorophyll content with level of irradiance. Use of a diving pulse amplitude modulated Xuorometer revealed that the zooxanthellae of R. rhodostoma eVectively disperse excess light energy by expressing significantly higher values of non-photochemical quenching and maximum excitation pressure on photosystem II when experimentally exposed to high light (HL) versus low light (LL). Host corallimorpharian tissues mediated this response by shielding the algal symbionts from high irradiance. The endoderm of host tentacles thickened significantly and microalgal cells were located further from the mesoglea in HL than in LL. The clades of zooxanthellae hosted by the corallimorpharians also varied with depth. In shallow water, all sampled individuals hosted clade C zooxanthellae, while in deep water the majority hosted clade D. The photosynthetic output of individuals of R. rhodostoma was less aVected by HL than was that of a stony coral examined. When exposed to both high temperature (HT) and HL, individuals of R. rhodostoma reduced their maximum quantum yield, but not when exposed to HL at low temperature (LT). In contrast, colonies of the scleractinian coral Favia favus reduced their photosynthetic

output when exposed to HL in both temperature regimes. After 2 weeks of HT stress, R. rhodostoma polyps appeared to bleach completely but reestablished their zooxanthella populations upon return to ambient temperature. We conclude that mechanisms of photoadaptation to high irradiance employed by both the endosymbiotic zooxanthellae and host corallimorpharians may explain in part the abundance of R. rhodostoma on some shallow reef Xats. The ability to survive for weeks at HT while bleached also may allow corallimorpharians to repopulate shallow reef areas where scleractinians have been killed by thermal stress. LaJeunesse, TC, Loh, WKW, van Woesik, R, Hoegh-Guldberg, O, Schmidt, GW and Fitt, WK. (2003) Low symbiont diversity in southern Great Barrier Reef corals, relative to those of the Caribbean. Limnology and Oceanography 48, 2046-2054. The specific identity of endosymbiotic dinoflagellates (Symbiodinium spp.) from most zooxanthellate corals is unknown. In a survey of symbiotic cnidarians from the southern Great Barrier Reef (GBR), 23 symbiont types were identified from 86 host species representing 40 genera. A majority (.85%) of these symbionts belong to a single phylogenetic clade or subgenus (‘‘C’’) composed of closely related (as assessed by sequence data from the internal transcribed spacer region and the ribosomal large subunit gene), yet ecologically and physiologically distinct, types. A few prevalent symbiont types, or generalists, dominate the coral community of the southern GBR, whereas many rare and/or specific symbionts, or specialists, are found uniquely within certain host taxa. The comparison of symbiont diversity between southern GBR and Caribbean reefs shows an inverse relationship between coral diversity and symbiont diversity, perhaps as a consequence of more-rapid diversification of Caribbean symbionts. Among clade C types, generalists C1 and C3 are common to both Caribbean and southern GBR symbiont assemblages, whereas the rest are regionally endemic. Possibly because of environmental changes in the Caribbean after geographic isolation through the Quaternary period, a high proportion of Caribbean fauna associate with symbiont taxa from two other distantly related Symbiodinium clades (A and B) that rarely occur in Pacific hosts. The resilience of Porites spp. And the resistance of Montipora digitata to thermal stress and bleaching are partially explained by their association with a thermally tolerant symbiont type, whereas the indiscriminant widespread bleaching and death among certain Pacific corals, during El Niño Southern Oscillation events, are influenced by associations with symbionts possessing higher sensitivity to thermal stress. LaJeunesse, TC, Thornhill, DJ, Cox, EF, Stanton, FG, Fitt, WK and Schmidt, GW. (2004) High diversity and host specificity observed among symbiotic dinoflagellates in reef coral communities from Hawaii. Coral Reefs 23, 596–603. The Hawaiian Islands represent one of the most geographically remote locations in the Indo-Pacific, and are a refuge for rare, endemic life. The diversity of symbiotic dinoflagellates (Symbiodinium sp.) inhabiting zooxanthellate corals and other symbiotic cnidarians from the High Islands region was surveyed. From the 18 host genera examined, there were 20 genetically distinct symbiont types (17 in clade C, 1 in clade A, 1 in clade B, and 1 in clade D) distinguished by internal transcribed spacer region 2 sequences. Most ‘‘types’’ were found to associate with a particular host genus or species and nearly half of them have not been identified in surveys of

Western and Eastern Pacific hosts. A clear dominant generalist symbiont is lacking among Hawaiian cnidarians. This is in marked contrast with the symbiont community structures of the western Pacific and Caribbean, which are dominated by a few prevalent generalist symbionts inhabiting numerous host taxa. Geographic isolation, low host diversity, and a high proportion of coral species that directly transmit their symbionts from generation to generation are implicated in the formation of a coral reef community exhibiting high symbiont diversity and specificity. LaJeunesse, TC, Reyes-Bonilla, H and Warner, ME. (2007) Spring “bleaching” among Pocillopora in the Sea of Cortez, Eastern Pacific. Coral Reefs 26, 265–270. A mild bleaching event was observed among Pocillopora spp. in the southern Gulf of California in the spring of 2006. Uniform bleaching occurred in numerous colonies on the upper portions of their branches. Most (»90%) colonies that exhibited bleaching contained a species of endosymbiotic dinoflagellate, Symbiodinium C1b-c, which differed from the Symbiodinium D1 found inhabiting most unbleached colonies. Analysis of chlorophyll fluorescence, indicated a decline in photosystem II photochemical activity, especially among colonies populated with C1b-c. By early August, most aVected colonies had recovered their normal pigmentation and fluorescence values were once again high for all colonies. No mortality was observed among tagged bleached colonies nor did symbiont species composition change during recovery. This unusual episode of bleaching did not appear to be a response to thermal stress, but may have been triggered by high levels of solar radiation during a period of unseasonally high water clarity in the early spring. Lam, KKY. (2003) Coral recruitment onto an experimental pulverised fuel ash-concrete artificial reef. Marine Pollution Bulletin 46, 642-653. An experimental artificial reef was deployed in December 1993 at Hoi Ha Wan Marine Park, Hong Kong. This is the first study documenting natural scleractinian coral recruitment onto a stabilised pulverised fuel ash (PFA)–concrete artificial reef. Visible recruits were first recorded 9–10 months after the placement of reef blocks, i.e., in the autumn of 1994. Two scleractinians, Oulastrea crispata and Culicia japonica, were recruited. The recruit density of the former was much greater than the latter. The spatial recruitment pattern of the corals was observed to be affected by the orientation of the attaching surface. O. crispata settled predominantly on the undersides of the reef blocks. There was an edge effect on O. crispata recruitment. C. japonica, however, had a preference for exposed surfaces. O. crispata did not show a preference for block composition whereas C. japonica favoured blocks with high (75% by volume) PFA levels. This shows that PFA–concrete is a potential substratum for artificial reef construction, especially when such reefs aim at rehabilitating corals.

Lan, C-H and Hsui, C-Y. (2005) The deployment of artificial reef ecosystem: Modelling, simulation and application. Simulation Modelling Practice and Theory. Artificial reefs (ARs) have been demonstrated to be a potential tool for the restoration of marine habitat and to retard the stress of coral reefs in recent decades. For most engineers, the deployment of an artificial reef ecosystem (ARE) is based on their experience, but what is the effective deployment to produce the higher abundance and to enrich biomass under the finite budget is seldom mentioned. In fact, how to deploy artificial reefs into the selected site and construct an effective ARE is the most challenge to ecological engineers. The main objective of this research is to construct a mathematical model called the deployment of artificial reef communities (DARCs) model, and then to provide a concrete approach for deploying an ARE under the finite budget. The constructing spirit of DARCs model is based on the observations of biologists that a higher complexity of habitat system will increase the species diversity and biomass. The fractal dimension (FD) is applied as an assessing indicator for the habitat system complexity of ARE in this study. In addition, a step-by-step algorithm for achieving the maximal fractal dimension of an ARE through the determinations of the number of deployed artificial reef communities (ARCs), the distance of adjacent ARCs, and the spatial configuration of ARCs in an ARE is presented. A computerized tool written by Fortran program to conduct the design of an ARE as well as perform the simulated analyses is completely proposed, and a case study is followed. The simulation-based analyses of changing budget reveal that the FD of a given region will be constrained regardless of increasing budget. This phenomenon shows that the more deployment of ARCs may lead to waste capital. Finally, a decision support system called DSSforARE is established for the quick design of an AR ecosystem under the consideration of conservation policy. Langdon, C, Broecker, WS, Hammond, DE, Glenn, E, Fitzsimmons, K, Nelson, SG, Peng, T-H, Hajdas, I and Bonani, G. (2003) Effect of eleveated CO2 on the community metabolism of an experimental coral reef. Global Biogeochemical Cycles 17, 1011. The effect of elevated pCO2 on the metabolism of a coral reef community dominated by macroalgae has been investigated utilizing the large 2650 m3 coral reef mesocosm at the Biosphere-2 facility near Tucson, Arizona. The carbonate chemistry of the water was manipulated to simulate present-day and a doubled CO2 future condition. Each experiment consisted of a one-two month preconditioning period followed by a seven-nine day observational period. The pCO2 was 404 ± 63 µatm during the present-day pCO2 experiment and 658 ± 59 µatm during the elevated pCO2 experiment. Nutrient levels were low and typical of natural reefs waters (NO3

- 0.5-0.9 µM, NH4

+ 0.4 µM, PO43- 0.07-0.09 µM). The temperature and salinity of the

water were held constant at 26.5 ± 0.2°C and 34.4 ± 0.2 ppt. Photosynthetically available irradiance was 10 ± 2 during the present-day experiment and 7.4 ± 0.5 mol photons m-2 d-1 during the elevated pCO2 experiment. The primary producer biomass in the mesocosm was dominated by four species of macroalgae: Haptilon cubense, Amphiroa fragillisima, Gelidiopsis intricata, and Chondria dasyphylla. Algal biomass was 10.4 mol C m-2 during the present-day and 8.7 mol C m-2 and during the elevated pCO2 experiments. As previously observed, the increase in pCO2 resulted in a decrease in calcification from 0.041 ± 0.007 to 0.006 ± 0.003 mol CaCO3 m

-2 d-1.

Net community production (NCP) and dark respiration did not change in response to elevated pCO2. Light respiration measured by a new radiocarbon isotope dilution method exceeded dark respiration by a factor of 1.2 ± 0.3 to 2.1 ± 0.4 on a daily basis and by 2.2 ± 0.6 to 3.9 ± 0.8 on an hourly basis. The 1.8-fold increase with increasing pCO2 indicates that the enhanced respiration in the light was not due to photorespiration. Gross production (GPP) computed as the sum of NCP plus daily respiration (light + dark) increased significantly (0.24 ± 0.03 vs. 0.32 ± 0.04 mol C m-2 d-1). However, the conventional calculation of GPP based on the assumption that respiration in the light proceeds at the same rate as the dark underestimated the true rate of GPP by 41-100% and completely missed the increased rate of carbon cycling due to elevated pCO2. We conclude that under natural, undisturbed, nutrient-limited conditions elevated CO2 depresses calcification, stimulates the rate of turnover of organic carbon, particularly in the light, but has no effect on net organic production. The hypothesis that an increase pCO2 would produce an increase in net production that would counterbalance the effect of decreasing saturation state on calcification is not supported by these data. Langmead, O and Sheppard, C. (2004) Coral reef community dynamics and disturbance: A simulation model. Ecological Modelling 175, 271-290. A spatially explicit model of coral community dynamics has been developed, based around a cellular automaton, with additional processes impacting upon it such as recruitment and disturbance. It represents a homogeneous plot on a Caribbean fore-reef slope with 10 coral species. Complexity that arises from species behaviour is shown to be realistic, generated by simple, repeated actions of the base units (coral polyps). The spatial resolution used here is high, and allows detailed demographic dynamics to be examined. Input parameters were based on values extracted from the literature for all processes (except disturbance, which is examined here). Natural background disturbance processes were explored in order to understand their importance in structuring coral communities and populations. Initially two disturbance parameters were investigated: spatial extent (proportion of plot affected) and size of disturbed patches. Both were found to be equally important in driving coral community structure, and subsequent diversity. The latter also influenced coral population size structure by altering mortality regimes. A power law model was introduced to distribute the total area disturbed into patches; its parameters were tuned using emergent size structure of the coral Agaricia spp. Differential mortality for species was also included, based on colony size and tissue regeneration rates. Model performance was assessed by running simulations with different levels of background disturbance, and resulting population size structures of each species were compared to field observations. Seven species compared extremely well, while accuracy of a further two could not be quantified due to a lack of suitable field data. Only one species, Siderastrea siderea, had unrealistic population size structures, which is discussed with respect to the life history of this species. Applications of this model include predicting communities under changing disturbance regimes, and colonisation and recovery processes following acute disturbances.

Ledlie, MH, Graham, NAJ, Bythell, JC, Wilson, SK, Jennings, S, Polunin, NVC and Hardcastle, J. (2007) Phase shifts and the role of herbivory in the resilience of coral reefs. Coral Reefs 26, 641–653. Cousin Island marine reserve (Seychelles) has been an effectively protected no-take marine protected area (MPA) since 1968 and was shown in 1994 to support a healthy herbivorous fish assemblage. In 1998 Cousin Island reefs suffered extensive coral mortality following a coral bleaching event, and a phase shift from coral to algal dominance ensued. By 2005 mean coral cover was <1%, structural complexity had fallen and there had been a substantial increase in macroalgal cover, up to 40% in some areas. No clear trends were apparent in the overall numerical abundance and biomass of herbivorous fishes between 1994 and 2005, although smaller individuals became relatively scarce, most likely due to the loss of reef structure. Analysis of the feeding habits of six abundant and representative herbivorous fish species around Cousin Island in 2006 demonstrated that epilithic algae were the preferred food resource of all species and that macroalgae were avoided. Given the current dominance of macroalgae and the apparent absence of macroalgal consumers, it is suggested that the increasing abundance of macroalgae is reducing the probability of the system reverting to a coral dominated state. Lee, SC. (2006) Habitat complexity and consumer-mediated positive feedbacks on a Caribbean coral reef. Oikos 112, 442-/447. Theoretical and empirical evidence suggest that positive feedbacks can increase resilience in ecological communities. On Caribbean coral reefs, there have been striking shifts from physically complex communities with high coral cover to relatively homogenous communities dominated by macroalgae, which have persisted for decades. However, little is known about positive feedbacks that may maintain coral reef community states. Here, I explore a potential consumer-mediated feedback on a Jamaican reef by examining how grazing by a keystone herbivore (Diadema antillarum) is enhanced by physical structure, which offer refugia from predation. Surveys revealed that habitat complexity and Diadema density were positively related. Increasing habitat complexity by adding physical structure significantly decreased macroalgal cover and increased the proportion of urchins in algal habitats in field manipulations. Experimental increases in urchin density also decreased macroalgal cover, but did not affect the proportion of urchins in algal habitats. These results suggest that the low habitat complexity of macroalgal-dominated reefs may inhibit an urchin-mediated shift to coral dominance and that positive feedbacks must be considered in reef restoration efforts. Leggat, W, Ainsworth, TD, Dove, S and Hoegh-Guldberg, O. (2006) Aerial exposure influences bleaching patterns. Coral Reefs 25, 452. On the 17th December 2005 Acropora millepora and other reef flat corals at Heron Island (23.44S, 151.91E) were found to have bleached on the west-north-west face of aerially exposed branches (Fig. 1). Visual surveys on the 15th and 16th had not detected any bleaching signs. In the preceding 3 days, water temperatures reached a maximum of 32.0°C on the 15th December and exceeded 31.0°C for only 2.5 h. Past studies suggest this alone is not sufficient to cause bleaching. Analysis of wind speed and direction for the morning of the 17th showed that the low tide at 03:00 hours

coincided with 4 h of high winds from the west-north-west. Peak gusts (57 km h-1) were within the highest 2.5% of all reading from the previous 6 years. Analysis of the red absorbing photosynthetic pigments (absorptivity) and dark-adapted fluorescent yield (Ralph et al. 2005) demonstrated a 65% loss of photosynthetic pigments and a 40% decrease in yield on the west-north-west face of exposed corals (Fig. 2) compared to the eastern side. Lesser, MP. (2004) Experimental biology of coral reef ecosystems. Journal of Experimental Marine Biology and Ecology 300, 217-252. Coral reef ecosystems are at the crossroads. While significant gaps still exist in our understanding of how ‘‘normal’’ reefs work, unprecedented changes in coral reef systems have forced the research community to change its focus from basic research to understand how one of the most diverse ecosystems in the world works to basic research with strong applied implications to alleviate damage, save, or restore coral reef ecosystems. A wide range of stressors on local, regional, and global spatial scales including over fishing, diseases, large-scale disturbance events, global climate change (e.g., ozone depletion, global warming), and over population have all contributed to declines in coral cover or phase shifts in community structure on time scales never observed before. Many of these changes are directly or indirectly related to anthropogenically induced changes in the global support network that affects all ecosystems. This review focuses on some recent advances in the experimental biology of coral reef ecosystems, and in particular scleractinian corals, at all levels of biological organization. Many of the areas of interest and techniques discussed reflect a progression of technological advances in biology and ecology but have found unique and timely application in the field of experimental coral reef biology. The review, by nature, will not be exhaustive and reflects the author’s interests to a large degree. Because of the voluminous literature available, an attempt has been made to capture the essential elements and references for each topic discussed. Lesser, MP and Farrell, JH. (2004) Exposure to solar radiation increases damage to both host tissues and algal symbionts of corals during thermal stress. Coral Reefs 23, 367-377. Elevated seawater temperatures have long been accepted as the principal stressor causing the loss of symbiotic algae in corals and other invertebrates with algal symbionts (i.e., ‘‘bleaching’’). A secondary factor associated with coral bleaching is solar irradiance, both its visible (PAR: 400–700 nm) and ultraviolet (UVR: 290–400 nm) portions of the spectrum. Here we examined the synergistic role of solar radiation on thermally induced stress and subsequent bleaching in a common Caribbean coral, Montastraea faveolata. Active fluorescent measurements show that steady-state quantum yields of photosystem II (PSII) fluorescence in the zooxanthellae are markedly depressed when exposed to high solar radiation and elevated temperatures, and the concentration of D1 protein is significantly lower in high light when compared to low light treatments under the same thermal stress. Both photosynthetic pigments and mycosporine-like amino acids (MAAs) are also depressed after experimental exposure to high solar radiation and thermal stress. Host DNA damage is exacerbated under high light conditions and is correlated with the expression of the cell cycle gene p 53, a cellular gatekeeper that modulates the

fate of damaged cells between DNA repair processes and apoptotic pathways. These markers of cellular stress in the host and zooxanthellae have in common their response to the enhanced production of reactive oxygen species during exposure to high irradiances of solar radiation and elevated temperatures. Taking these results and previously published data into consideration, we conclude that thermal stress during exposure to high irradiances of solar radiation, or irradiances higher than the current photoacclimatization state, causes damage to both photochemistry and carbon fixation at the same time in zooxanthellae, while DNA damage, apoptosis, or necrosis are occurring in the host tissues of symbiotic cnidarians. Lewis, CL and Coffroth, MA. (2004) The acquisition of exogenous algal symbionts by an octocoral after bleaching. Science 304, 1490-1492. Episodes of coral bleaching (loss of the symbiotic dinoflagellates) and coral mortality have occurred with increasing frequency over the past two decades. Although some corals recover from bleaching events, the source of the repopulating symbionts is unknown. Here we show that after bleaching, the adult octocoral Briareum sp. acquire dinoflagellate symbionts (Symbiodinium sp.) from the environment. Uptake of exogenous symbionts provides a mechanism for response to changes in the environment and resilience in the symbiosis. Lien, Y-T, Nakano, Y, Plathong, S, Fukami, H, Wang, J-T and Chen, CA. (2007) Occurrence of the putatively heat-tolerant Symbiodinium phylotype D in high-latitudinal outlying coral communities. Coral Reefs 26, 35–44. Biogeographic investigations have suggested that coral-symbiont associations can adapt to higher temperatures by hosting a heat-tolerant Symbiodinium, phylotype D. It is hypothesized that phylotype D is absent in high latitudes due to its heattolerant characteristics. In this study, this hypothesis was tested by examining the symbiont diversity in a scleractinian coral, Oulastrea crispata, throughout its entire latitudinal distribution range in the West Pacific. Molecular phylotyping of the 5’-end of the nuclear large subunit of ribosomal DNA (lsu rDNA) indicated that phylotype D was the dominant Symbiodinium in O. crispata from the tropical reefs to the marginal nonreefal coral communities. Several colonies of tropical populations were associated with phylotype C, either alone or simultaneously with phylotype D. Analysis of the polymerase chain reaction products using singlestrand conformation polymorphism (SSCP) detected relatively low densities of phylotype C in most of the O. crispata colonies surveyed. These results provide evidence for the occurrence of phylotype D in coldwater outlying coral communities. The dominant occurrence of phylotype C in some O. crispata colonies on tropical reefs and the relatively low densities of phylotype C identiWed by SSCP in subtropical and temperate populations show that the dominant symbiont type can vary in this coral species and that multiple symbionts can co-occur in the same host.

Lindahl, U, Ohman, MC and Schelten, CK. (2001) The 1997/1998 mass mortality of corals: Effects on fish communities on a Tanzanian coral reef. Marine Pollution Bulletin 42, 127-131. The abnormally high surface temperatures in the world's oceans during 1997/1998 resulted in widespread coral bleaching and subsequent coral mortality. An experiment was performed to study the effects of this coral mortality as well as the influence of the structural complexity on fish communities on a Tanzanian coral reef. Changes in fish communities were investigated on plots of transplanted corals after 88% of these corals had died. A distinct shift in fish community composition was found, although diversity was not affected. Fish abundance rose by 39% mostly due to an increase in herbivores, which seemed to benefit from enhanced algal growth on the dead corals. Fish abundance, species diversity and community composition were also strongly influenced by the structural complexity provided by the live and dead corals. This suggests that a coral reef can support abundant and diverse fish populations also after the corals have died as long as the reef structure is sustained. Lindén, O, Souter, D, Wilhelmsson, D and Obura, D (eds.) (2002) Coral reef degradation in the Indian Ocean: Status report 2002. CORDIO: Kalmar (Sweden), 284 pp. Since the 1998 coral bleaching and mortality, which affected reefs throughout the tropics, a number of projects dealing with research, monitoring and management of coral reefs have been conducted. A number of these projects have been carried out under the CORDIO Program in the Indian Ocean where coral reefs have been severely impacted by bleaching related mortality and other anthropogenic influences. This, CORDIO’s third status report, describes the condition of coral reefs throughout the central and western Indian Ocean and presents the results of several targeted research projects conducted within the last 18 months. The Recovery of Coral Communities Including Fish The covery of coral reefs is highly variable between sites in East Africa and South Asia. Generally, extensive mortality affected the islands of the northern and central Indian Ocean (Maldives, Seychelles, Aldabra, Chagos), the East African coast (Mozambique, Tanzania and Kenya), Sri Lanka and parts of India. Islands of the southern Indian Ocean (Reunion, Comoros, Mauritius, Madagascar) were not seriously affected. In most of the areas that suffered high coral mortality (i.e. more than 50%), recovery is slow, patchy or non-existent. In some areas, significant recovery appears to be underway. However, such areas are usually located far away from human influence or are well protected in marine reserves. It is obvious that in many places closer to human habitation other stresses from anthropogenic activities, such as destructive fishing, coral mining and sedimentation from land, continue to degrade the reefs, slowing down or preventing recovery from the 1998 bleaching event. TANZANIA Little recovery was observed on the reefs of Misali Island where 80% mortality was noted in 1998. Similar observations were made on Kwale and Mafia reefs where the cover of live coral remained low or continued to decline. Chumbe Island and the reefs off Zanzibar town were relatively unaffected by the 1998 bleaching event. However destructive fishing and outbreaks of Crown of Thorns starfish are now affecting several of these reefs. Fish populations showed significant changes and several

groups (especially invertivores, piscivores and plaktivores) decreased while other groups (particularly herbivores) increased. During 2001–02, harmful algal blooms and an unknown fungal coral disease have impacted reefs in northern Tanzania. These stresses may be climate-change related. Over-fishing, destructive fishing, pollution from human settlement and development, mining and shipping industry activities continue to cause degradation to the reefs of the country. KENYA The coral reefs were severely impacted in 1998, with bleaching and mortality levels of 80% and greater. Recovery of affected reefs to 2002 has been moderate and patchy. Reefs with high coral diversity and cover that were strongly impacted in 1998 have recovered to less than 50% of pre-bleaching coral cover. Some high diversity reefs that were protected from bleaching impacts have remained healthy. Degraded reefs outside of Marine Protected Areas that were strongly impacted by the El Niño have generally recovered to 50–100% of prebleaching coral cover. In 2001–02, multiple threats have impacted Kenyan reefs that may be climate-change related, including harmful algal blooms and an unknown fungal coral disease. Anthropogenic activities including over-fishing, destructive fishing, pollution, human settlement and development, mining and shipping industry activities continue to threaten East African reefs. MOZAMBIQUE In the two years after 1998, reefs that had been severely affected by the bleaching showed slow or no recovery. Reefs in northern remote areas and in protected areas showed increasing coral cover. In February 2000, severe floods affected the country. These floods brought large quantities of sediment to the reefs and caused significant reduction of the remaining coral cover: from 20 and 5% to 10 and 0.5% respectively for hard and soft corals in the Xai-Xai lagoon. MADAGASCAR Information is limited to a few areas in the northwest and southwest. In the northwest the corals were relatively unaffected by the bleaching and the ecosystem was in good condition. In the southwest the reefs were little impacted and recovery is underway. SEYCHELLES The reefs of the inner granitic islands suffered extensive and system-wide degradation in 1998 and hard coral abundance and diversity are now well below pre-1998 levels. However, small but significant increases in coral cover and diversity have been noted over the past 15 months. Recovery may be underway in the near-to midterm. The rate at which local coral communities will recover will be largely dependent on the environmental conditions encountered by newly settled coral recruits. REMOTE REEFS OF THE CENTRAL INDIAN OCEAN The cover of live coral at Aldabra has changed little between 1999 and 2001 indicating that recovery is a slow process even in remote areas beyond the reach of anthropogenic influences. Nevertheless, recovery of the reef at Aldabra is unlikely to be limited by the influx of new corals as the average number of recruits per m2 ranged between 4 and 7 with the greatest number recorded at approximately 10 m depth. The impacts of bleaching related coral mortality on Aldabra’s fish communities was difficult to discern. The abundance of fish at Aldabra varied both spatially and temporally but the diversity of fish species has remained high with 287 species from 35 families being recorded in 1998 prior to the bleaching event, 211 species from 35 families in 1999 and 205 species from 40 families in 2001. Also, in Chagos, where most corals died in 1998, the recovery is very slow and is being hampered by the

breakdown of the dead coral thickets due to bio-erosion forming large areas of unstable substrates. MALDIVES The recruitment of corals in Maldives is well underway, but is patchy and does not resemble the pre-bleaching community structure suggesting that these reefs will experience a change in community structure. SRI LANKA The recovery of shallow reefs in Sri Lanka is variable between sites but has, in general, been slow and in some areas non-existent. Algae, tunicates and corallimorpharians largely dominate dead reefs. Fish populations have changed and several of the typical coral fish species have become rare or have disappeared. Destructive human activities such as coral mining, extensive fishing and collection of fish for the aquarium trade continue to degrade the reefs preventing recovery. REEFS OF THE GULF OF MANNAR AND NICOBAR AND ANDAMAN ISLANDS Recent reports from the Gulf of Mannar show variable results, which might indicate both significant recovery and that the impact in 1998 was patchy affecting certain areas while others did not suffer to the same extent. Recent surveys from the Nicobar and Andaman Islands show no signs of a mass mortality in 1998. Socio-Economic Effects Fish populations were affected by the bleaching both in terms of species composition and the total quantities of fish . However, more research will be needed to establish the detailed relationships and quantify the effects. Results from certain areas seriously affected by bleaching and subsequent coral mortality indicated an initial increase in catch figures caused by increased populations of herbivorous species that were able to utilize the overgrowth of fast-growing algae that colonised the reefs after the corals had died. It is difficult to directly link decreasing fish catches to the bleaching event. The general trend in most of the shallow reef areas in the region is one of declining stocks due to over-fishing and habitat destruction. However, the data from different studies carried out in the region are not conclusive. It should be pointed out that observations of impacts on fish populations through catch data are often unreliable, as catch statistics do not usually reflect the situation in the environment. If traditional sites yield fewer fish, fishermen usually move to other areas and/or target other species or use different gear. In addition, the catch per unit effort may appear to be stable or even increasing giving the impression that the fish stocks are healthy and under-utilized. However, increased catches are often the result of the use of more efficient fishing gear and boats and not at all a true reflection of the condition of fish stocks. The general trend in most areas in the region is that the fishing pressure is increasing. In addition, the official statistics often have little to do with the real catches, as much of what is caught is not recorded. What is obvious from several observations throughout the region is that there has been a shift in fish species composition in areas severely affected by the bleaching. Species such as butterfly fish and surgeon fish are decreasing in numbers in areas where the coral has died while some other species such as rabbit fish as well as small pelagic species such as clupeids appears to increase, at least in the short-term. Results from several areas indicate that, as long as the reef structure remains intact, the reefs are still able to provide protection and function as nursery areas for juvenile fish and other invertebrates. However, once the structure of the reefs collapses, these functions are likely to be lost. Observations from Chagos indicate that extensive areas of reef that died in 1998 have been eroded to rubble and are now forming a substrate too unstable to allow settlement of coral larvae.

The impacts of coral bleaching on tourism have also been variable across the western Indian Ocean region. In saturated tourist destinations such as the Maldives, only slight impacts have been observed because the industry was able to compensate for losses incurred in dive and other reef-related tourism sectors with gains in other areas such as the honeymoon market. In less popular destinations the impacts were more severe. According to surveys carried out in Sri Lanka in 2000–2001, the snorkelling and glass bottom boat industry have declined as a result of the mass bleaching of corals in 1998. Detailed investigations of the impacts of bleaching on tourism in Seychelles, Kenya and Zanzibar indicated that the possibility to enjoy healthy reefs was an important component of the holidays of tourists at these destinations. On average, tourists were ‘willing to pay’ US$ 98.7 extra per holiday in the Seychelles, US$ 87.7 extra in Zanzibar and US$ 59.0 extra in Kenya in order to experience high quality reefs indicating that healthy reefs are important assets for tourism in the coastal regions of the Indian Ocean and that coral reef degradation could potentially cause a significant decline in tourism in these areas. Moreover, continued degradation of coral reefs will have a significant impact on the economies of those countries in the western Indian Ocean. Based on tourists willingness to pay, the number of tourists visiting Seychelles, Mombasa and Zanzibar and their interest in the marine environment, the economic losses were estimated to be US$ 71.5 million for the Seychelles, US$ 47.2 million for Mombasa and US$ 39.9 million for Zanzibar in net present values over a 20 year period with a 10% discount rate. Public Awareness and Alternative Livelihoods In addition to ecological and socio-economic research in the Indian Ocean, CORDIO, in collaboration with other organizations, is involved in projects to inform the public, staff of governmental agencies and politicians of the need for better management of costal marine ecosystems including coral reefs. For example, during 2001, CORDIO supported an awareness project titled ‘A tomorrow for our reefs’ implemented by IUCN Sri Lanka. In addition, on the Tuticorin Coast of India, and with WWF as a lead partner in Kiunga, Kenya, community based reef restoration has been initiated. Coral fragments are cultivated and transplanted to the reefs by the local community members thus restoring parts of the reefs and creating an understanding of the value and importance of the corals among the local communities in the area. Raising awareness and building community capacity to participate in management of reefs and to improve local livelihoods is one goal of a CORDIO project in Diani, Kenya. Here, monitoring of reef fish catch, underwater fish populations and socioeconomic factors is being undertaken by fishermen and community members in partnership with scientists. While the data contributes to knowledge on reef health and status of the fishery, the process of monitoring has a strong educational and empowerment component, encouraging participating community groups to take a lead in caring for the coral reefs they depend on. The project is serving as a model for other community-based projects in East Africa working with coral reefs. The results of a number of studies show the necessity to reduce the pressure from both legal and illegal human activities on the reefs, through better management and law enforcement. A high and growing population pressure characterizes many coastal areas. In order relieve the pressures on the ecosystem, there is a great need for alternative income sources for the people living in areas where the ecosystem is degrading. In Sri Lanka, a project has been started to assess the experiences of previous efforts and initiatives as well as experiences from other regions to generate alternative income sources. The outcome of this project will result in

recommendations to relevant authorities in Sri Lanka, and for the basis for small-scale pilot projects under the CORDIO program starting 2003. The Way Forward The scientific assessments carried out under the CORDIO program are done by local scientists and experts in 11 countries around the Indian Ocean. Hence, CORDIO is a rare example of a locally driven, regional initiative that provides valuable data not only for the countries of the region but also for the world at large. It is important that such networks are maintained, because the collection of long-term data series is the only process that enables changes in the state of the environment to be determined and the potential threats to coastal populations and societies to be mitigated. The CORDIO program will continue to support and coordinate ecological and socio-economic monitoring of reefs in the Indian Ocean region with the aim of studying the processes of coral reef recovery from the 1998 coral bleaching, and reporting the state of the coastal environments. The long-term information obtained from continued monitoring will be used to develop projects investigating new and improved management options in several pilot areas in the region. This work will involve the regional nodes of CORDIO in Mombasa, Colombo and St. Denis. In addition, with a closer collaboration with the World Conservation Union (IUCN) agreements will be made between CORDIO regional offices and both IUCN national and regional marine programs to develop this type of activities. Lirman, D, Manzello, D and Macia, S. (2002) Back from the dead: the resilience of Siderastrea radians to severe stress. Coral Reefs 21, 291–292. Siderastrea radians is considered one of the most stress-tolerant corals species in the Caribbean, often found in highly disturbed locations (Lewis 1989). In Biscayne Bay, Florida, a coastal lagoon adjacent to the city of Miami, S. radians is the dominant coral species, reaching densities of up to 68 colonies m–2. This shallow basin (1–3 m) experiences high sedimentation, rapidly fluctuating salinity, and temperature extremes that make it a marginal environment for coral growth. Little, AF, van Oppen, MJ and Willis, BL. (2004) Flexibility in algal endosymbioses shapes growth in reef corals. Science 304, 1492-1494. The relation between corals and their algal endosymbionts has been a key to the success of scleractinian (stony) corals as modern reef-builders, but little is known about early stages in the establishment of the symbiosis. Here, we show that initial uptake of zooxanthellae by juvenile corals during natural infection is nonspecific (a potentially adaptive trait); the association is flexible and characterized by a change in (dominant) zooxanthella strains over time; and growth rates of experimentally infected coral holobionts are partly contingent on the zooxanthella strain harbored, with clade C-infected juveniles growing two to three times as fast as those infected with clade D. Liu, G, Strong, AE and Skirving, W. (2003) Remote sensing of sea surface temperatures during 2002 Barrier Reef coral bleaching. EOS 84, 137-144. Early in 2002, satellites of the U.S.National Oceanic and Atmospheric Administration (NOAA) detected anomalously high sea surface temperatures (SST) developing in the western Coral Sea, midway along Australia’s Great Barrier Reef (GBR).This was

the beginning of what was to become the most significant GBR coral bleaching event on record [Wilkinson, 2002]. During this time, NOAA’s National Environmental Satellite,Data,and Information Service (NESDIS) provided satellite data as part of ongoing collaborative work on coral reef health with the Australian Institute of Marine Science (AIMS) and the Great Barrier Reef Marine Park Authority (GBRMPA).These data proved invaluable to AIMS and GBRMPA as they monitored and assessed the development and evolution of SSTs throughout the austral summer, enabling them to keep stakeholders,government, and the general public informed and up to date. As anomalous SSTs continued to increase off the coast of Queensland, NOAA, AIMS, and GBRMPA issued a press release on 24 January 2002 on the developing bleaching event engulfing much of the mid- and northern GBR, which later proved to be the most extensive bleaching event on record for that region. Bleaching occurs when there is widespread loss of pigment from coral, mainly due to the expulsion of symbiotic algae [Yonge and Nicholls, 1931]. The algae are usually expelled in times of stress, often caused by sea surface temperatures which are higher than the coral colony’s tolerance level. This may be as little as 1 to 2°C above the mean monthly summer values [Berkelmans and Willis, 1999;Reaser et al., 2000]. The key coral bleaching products developed by the satellite monitoring and prediction component of NOAA’s Coral Reef Watch (CRW) program include: SST coral bleaching “HotSpot” anomaly and “Degree Heating Week”(DHW) charts. In this article, these products will be used to describe the evolution of the anomalous SST event that caused the 2002 GBR bleaching event, which will then be contrasted with the 1998 bleaching event. The latter was the largest for the GBR region prior to the 2002 event (R.Berkelmans, Austral. Inst.of Marine Science, unpublished data, 2002). Loch, K, Loch, W and Schuhmacher, H. (2002) Coral recruitment and regeneration on a Maldivian reef 21 months after the coral bleaching event of 1998. Marine Ecology 23, 219-236. This paper describes the quantitative inventory of stony corals on a Maldivian reef after the bleaching event of 1998. The detailed data, collected in March 1999 and March 2000, comprise survival, new recruitment and regenerates. They were obtained in 6 transects laid out randomly on the reef flat, on 22 Acropora tables on 6 sites at the reef edge and on 39 Porites lobata blocks and 1 Diploastrea heliopora colony. The present cover of living zooxanthellate corals is reduced to ca. 2 – 5 % of its previous state. Acroporidae and Pocilloporidae were practically wiped out, while Poritidae survived partly and Agariciidae (esp. Pavona) are now the dominant group. New settlements on dead Acropora tables were mainly Agariciidae, followed by Acroporidae and Pocilloporidae. Regenerates on Porites were pronounced and the apparent yearly increase in mass was about threefold that of Diploastrea, which is 3-4mm per year. The influence on reef ecology in terms of coral substrate and species, possible sources of larvae, change of guilds in reef builders, other species and the prospect for further development of the reef, with respect to growth versus erosion, is discussed.

Loch, K, Loch, W, Schuhmacher, H and See, WR. (2004) Coral recruitment and regeneration on a Maldivian reef four years after the coral bleaching even of 1998. Part 2: 2001-2002. Marine Ecology 25, 145-154. This paper is the follow-up study to a quantitative inventory of stony corals on a Maldivian reef after the bleaching event of 1998 [Loch, K., W. Loch, H. Schuhmacher & W. R. See, 2002; P.S.Z.N.: Marine Ecology, 23(3): 219–236] covering the years 2001 and 2002, thereby extending the observation period from 1999 to 2002. Data collection was performed as previously in transects on the reef flat, Acropora tables at the reef rim, Porites lobata blocks and one Diploastrea heliopora colony. While between 2000 and 2001 the transects showed a noticeable increase of living scleractinian colonies of 6.1 per m2 of reef surface per year, this had slowed to 0.7 per m2 per year by 2002. Likewise, the colonisation of dead Acropora tables had changed from a yearly increase of 14.4 colonies per m2 of table surface between 2000 and 2001 to a decrease of 9.4 between 2001 and 2002. The main constituent of the regeneration in transects as well as on Acropora tables was Pavona (Agariciidae), whereas Pocilloporidae and Acroporidae were almost completely missing. A new immigrant was Tubastraea micranthus. Continuous growth of regenerates on Porites presented a columnar form, and the growth rate of D. heliopora was reduced between 2001 and 2002. The influence on reef ecology in terms of coral substrate, species differentiation, growth rate changes, interaction with other organisms and the overall prospect of the reef development on a longer timescale is outlined. Lombardi, MR, Lesser, MP and Gorbunov, MY. (2000) Fast repetition rate (FRR) fluorometry: Variability of chlorophyll a fluorescence yields in colonies of the corals, Montastrea faveolata (w.) and Diploria labyrinthiformes (h.) recovering from bleaching. Journal of Experimental Marine Biology and Ecology 252, 75-84. Recently, an underwater version of a fast repetition rate fluorometer (FRRF) was developed for the non-destructive study of fluorescence yields in benthic photoautotrophs.We used an FRRF to study bleached colonies of the corals, Montastraea faveolata and Diploria labyrinthiformes at sites surrounding Lee Stocking Island, Exuma, Bahamas, to assess their recovery from bleaching (~1 year after the initial bleaching event) induced by elevated temperatures. The steady state 9 quantum yields of chlorophyll a fluorescence (∆F’/F’m) from photosystem II (PSII) within coral colonies were separated into three categories representing visibly distinct degrees of bleaching ranging from no bleaching to completely bleached areas. Differences in (∆F’/F’m) were significantly different from bleached to unbleached regions within colonies. Dark, unbleached regions within colonies exhibited significantly higher (∆F’/F’m) values (0.438±0.019; mean±S.D.) when compared to lighter regions, and occupied a majority of the colonies’ surface area (46–73%). Bleached regions exhibited significantly lower (∆F’/F’m) (0.337±0.014) and covered only 7–25% of the colonies’ surface area. The observations from this study suggest that zooxanthellae in bleached regions of a colony exhibit reduced photosynthetic activity as long as one year after a bleaching event and that in situ fluorescence techniques such as FRRF are an effective means of studying coral responses and recovery from natural or anthropogenic stress in a non-destructive manner.

Lough, J. (1999) Sea surface temperatures on the Great Barrier Reef: A contribution to the study of coral bleaching. Great Barrier Reef Marine Park Authority: Townsville (Australia), 31 pp. Abstract not available. Loya, Y, Sakai, K, Yamazato, K, Nakano, H, Sambali, H and van Woesik, R. (2001) Coral bleaching: The winners and losers. Ecology Letters 4, 122-131. Sea surface temperatures were warmer throughout 1998 at Sesoko Island, Japan, than in the 10 preceding years. Temperatures peaked at 2.8 °C above average, resulting in extensive coral bleaching and subsequent coral mortality. Using random quadrat surveys, we quantitatively documented the coral community structure one year before and one year after the bleaching event. The 1998 bleaching event reduced coral species richness by 61% and reduced coral cover by 85%. Colony morphology affected bleaching vulnerability and subsequent coral mortality. Finely branched corals were most susceptible, while massive and encrusting colonies survived. Most heavily impacted were the branched Acropora and pocilloporid corals, some of which showed local extinction. We suggest two hypotheses whose synergistic effect may partially explain observed mortality patterns (i.e. preferential survival of thick-tissued species, and shape-dependent differences in colony mass transfer efficiency). A community-structural shift occurred on Okinawan reefs, resulting in an increase in the relative abundance of massive and encrusting coral species. Loya, Y. (2007) How to influence environmental decision makers? The case of Eilat (Red Sea) coral reefs. Journal of Experimental Marine Biology and Ecology 344, 35–53. In a recent overview of the literature, Rinkevich (2005b), hereafter ‘Rinkevich’, aims to critically examine some of the paradigms concerning the degradation of Eilat's coral reef. He selects 5 of the major challenging outcomes/points he identifies, to motivate critical evaluation of the published literature and future research activities. He further states that his synopsis may also serve as an important application tool for managers and policy makers (p.185). In view of the widely publicized and intense ongoing (7–8 years) scientific and public debate on the possible connection between intensive fish farms' activity in Eilat and the state of the reef (see Erez et al., 2000; Genin, 2000; Gordin, 2000; Lazar et al., 2000; Atkinson et al., 2001; Genin et al., 2001; Bongiorni et al., 2003; Loya and Kramarsky-Winter, 2003; Rinkevich et al., 2003; Wielgus et al., 2003, 2004; Loya, 2004a; Atkinson et al., 2004; Loya et al., 2004, 2005; Abelson et al., 2005; Rinkevich, 2005a; Israel Ministry of Environmental Protection, 2006), Rinkevich's synopsis might prove to be critical for the possibility of legal measures that may (or may not) be undertaken, in the near future, with respect to the fish farm industry. Thus, the fate of the fish farms awaits the verdict of the Israeli Supreme Court, to whom they appealed with a request to call off a Governmental ruling (from June 2005) to stop their activity within 3 years (for landmarks in the fish farms controversy, see Appendix A). Since Rinkevich's paper has been published in a respected scientific journal, it now constitutes a significant document whichmight influence decisionmakers involved in making balanced judgments about reef protection in the northern Gulf of Eilat. It is therefore important

to draw the reader's attention to a number of problems with Rinkevich's paper, which I believe substantially affect his main conclusions. These problems centre on biased presentation and interpretation of refereed and grey literature that he claims support his assertions, but which are flawed and therefore misleading. Rinkevich disputes in particular series of my publications dealing with Eilat's coral community structure and species diversity, from the early 70's to date. These publications are central to the 5 points, discussed by Rinkevich. The purpose of this paper is not to defend my earlier work, or my recent publications dealing with the state of Eilat's reef, but to call attention to the severe state of the reef, (contrasting Rinkevich's declarations) and the urgent need to protect it. Lugo-Fernández, A, Deslarzes, KJP, Price, JM, Boland, GS and Morin, MV. (2001) Inferring probable dispersal of Flower Garden Banks coral larvae (Gulf od Mexico) using observed and simulated drifter trajectories. Continental Shelf Research 21, 47-67. We investigated likely coral larvae dispersal in the Gulf of Mexico by means of a combination of satellite-tracked drifters and simulated currents. Drifter data were collected during spawning events in 1997 and 1998 at the Flower Garden Banks (FGB) and from the “Surface Current and Lagrangian Drift Program'' in 1993 (Niiler et al., Surface current and langrangian-drift program. Unpublished Report, U.S. Department of the Interior, Minerals Management Service, 1997, 10pp). Observed and simulated tracks showed that likely coral dispersal is driven by five circulation modes. Mode 1 is a cyclonic motion reaching the Texas shelf and possible cyclonic motion along the shelf toward Mexico. Mode 2 is an entrainment by offshore eddies causing likely larval transport into deep water (>200 m). Mode 3 consists of a continuous eastward movement that reaches 87.58W. Mode 4 consists of recirculating flows over or near the FGB. Mode 5 is a cross-basin transport that ends near the Florida Keys in 50-60 days. Modes 1 and 4 can bring larvae back to the FGB within 24-30 days after spawning creating conditions for self-seeding. Offshore eddies near the FGB, Modes 2 and 5, generate offshore transport that can carry coral larvae to reefs in Mexico and Florida. Only ~43% of the drifters leave the northwestern Gulf shelf, which implies that most coral larvae can remain on the shelf through Modes 1, 3, and 4. Mode 3 suggests eastward larval dispersal and under influence of theMississippi River plume where they can be adversely affected by low salinity, low temperature, and high sedimentation which limit recruitment on oil and gas platforms. Possible effects of hurricanes on larval dispersal include reduced travel time, considerable displacements, and physical damage by strong turbulence. Nineteen outer shelf banks were contacted by actual and simulated drifter tracks, but only 10 are suitable for coral larvae settlement given the substrate and crest depth. These 10 banks could represent a doubling of reef area relative to the FGB, i.e., 5.35km2 shallower than 55 m, if successfully colonized by coral larvae. Drifter tracks also contacted 129 major oil and gas fixed platforms. Potential larvae recruitment by these contacts could increase the reef area in the northwestern Gulf by 38% compared to the FGB, and support eastward propagation of corals. A majority of actual and simulated drifter contacts occurs within 40km of the FGB; contacts increased at 70km and beyond 150km from the FGB. This contact distribution suggest a high potential for self-seeding, and connectivity of Gulf reefs at long time scales.

Lyons, MM, Aas, P, Pakulski, JD, van Waasbergen, L, Miller, RV, Mitchell, DL and Jeffrey, WH. (1998) DNA damage induced by ultraviolet radiation in coral-reef microbial communities. Marine Biology 130, 537-543. Ultraviolet radiation (UVR) has been implicated in coral-bleaching processes and UVR may create stress to marine organisms by damage to DNA. Absorption of energy from UVB (280 to 320 nm) induces direct damage to DNA via cyclobutane pyrimidine dimer photoproduct-formation. We examined the extent of DNA damage created by UVR in coral reef microbial communities and whether the coral-surface microlayer (CSM) provides protection from UVR to the microorganisms found there. Diel patterns and depth profiles of UVR effects were examined in coral mucus (coral-surface microlayer, CSM) from Montastraea faveolata and Colpophyllia natans, and water-column samples of similar depths. UV-induced photodamage was determined using a radioimmunoassay specific for cyclobutane pyrimidine dimers (thymine dimers). Significant photo-damage was detected in water-column and CSM samples, although the level of damage in CSM samples was consistently lower than in water-column samples collected from the same depth, suggesting the presence of photoprotective mechanisms within the CSM. Diel patterns of photodamage were detected in both water-column and CSM samples, but peak damage occurred earlier in the day for the CSM samples, suggesting differences in damage and repair kinetics between the water column and CSM. The results suggest that microorganisms within the CSM are afforded some protection from UVR stress and that changes in the amount of DNA damage in these organisms may be an indicator of changing UVR stress to corals. Macintyre, IG, Glynn, PW and Hinds, F. (2005) Evidence of the role of Diadema antillarum in the promotion of coral settlement and survivorship. Coral Reefs 24, 273. In 1993, a seawall consisting of large limestone blocks was constructed off the south coast of Barbados to prevent beach erosion during major storms. In September 2004, we observed numerous mature Diadema antillarum on the leeward side of this wall (Fig.1a), in concentrations of up to 5/sq. m. These sea urchins had grazed and cleared some surface areas of filamentous and macroalgae, evidently promoting the growth of both crustose coralline algae and a variety of corals, including Diploria strigosa, Porites porites, Porites astreoides, and Siderastrea siderea. In marked contrast, the blocks on the seaward side of this wall showed no indication of D. antillarum, and the surface cover was dominated by filamentous algae with some scattered macroalgal growths, most notably Dictyota divaricata (Fig. 1b). The lack of D. antillarum on the seaward side is probably related to resuspended sediment and high bottom wave velocities in this exposed setting. Although there are several reports of the role of D. antillarum in the promotion of coral settlement (e.g., Hughes et al. 1987; Edmunds and Carpenter 2001), this observation of two sides of a seawall appears to provide an excellent one-site record of the enhancement of coral-community development by this sea urchin, which suffered mass mortality in 1982-1983, to the detriment of many coral communities in the Caribbean (Carpenter 1997).

Mantyka, CS and Bellwood, DR. (2007) Direct evaluation of macroalgal removal by herbivorous coral reef fishes. Coral Reefs 26, 435–442. Few studies have examined the relative functional impacts of individual herbivorous fish species on coral reef ecosystem processes in the Indo-Pacific. This study assessed the potential grazing impact of individual species within an inshore herbivorous reef fish assemblage on the central Great Barrier Reef (GBR), by determining which fish species were able to remove particular macroalgal species. Transplanted multiple-choice algal assays and remote stationary underwater digital video cameras were used to quantify the impact of local herbivorous reef fish species on 12 species of macroalgae. Macroalgal removal by the fishes was rapid. Within 3 h of exposure to herbivorous reef fishes there was significant evidence of intense grazing. After 12 h of exposure, 10 of the 12 macroalgal species had decreased to less than 15% of their original mass. Chlorodesmis fastigiata (Chlorophyta) and Galaxaura sp. (Rhodophyta) showed significantly less susceptibility to herbivorous reef fish grazing than all other macroalgae, even after 24 h exposure. Six herbivorous and/or nominally herbivorous reef Wsh species were identified as the dominant grazers of macroalgae: Siganus doliatus, Siganus canaliculatus, Chlorurus microrhinos, Hipposcarus longiceps, Scarus rivulatus and Pomacanthus sexstriatus. The siganid S. doliatus fed heavily on Hypnea sp., while S. canaliculatus fed intensively on Sargassum sp. Variation in macroalgal susceptibility was not clearly correlated with morphological and/or chemical defenses that have been previously suggested as deterrents against herbivory. Nevertheless, the results stress the potential importance of individual herbivorous reef fish species in removing macroalgae from coral reefs. Marshall, PA and Baird, AH. (2000) Bleaching of corals on the Great Barrier Reef: Differential susceptibilities among taxa. Coral Reefs 19, 155-163. Large-scale coral bleaching episodes are potentially major disturbances to coral reef systems, yet a definitive picture of variation in assemblage response and species susceptibilities is still being compiled. Here, we provide a detailed analysis of the bleaching response of 4160 coral colonies, representing 45 genera and 15 families, from two depths at four sites on reefs fringing inshore islands on the Great Barrier Reef. Six weeks after the onset of large-scale bleaching in 1998, between 11 and 83% of colonies along replicate transects were affected by bleaching, and mortality was 1 to 16%. There were significant differences in bleaching response between sites, depths and taxa. Cyphastrea, Turbinaria and Galaxea were relatively unaffected by bleaching, while most acroporids and pocilloporids were highly susceptible. The hydrocorals (Millepora spp.) were the most susceptible taxa, with 85% mortality. Spatial variation in assemblage response was linked to the taxonomic composition of reef sites and their bleaching history. We suggest, therefore, that much of the spatial variation in bleaching response was due to assemblage composition and thermal acclimation.

Marshall, P. (2003) Great Barrier Reef Coral Bleaching Response Program. Great Barrier Reef Marine Park Authority: Townsville (Australia), 15 pp. Large-scale coral bleaching events, driven by unusually warm sea temperatures, have now affected every major coral reef ecosystem on the planet. The effects of coral bleaching are pervasive, and potentially devastating to ecosystems and the people who depend upon them. The frequency and severity of these large-scale disturbances is predicted to increase as temperatures continue to warm under a global regime of climate change. In combination with the multitude of other stressors resulting from human activities, climate change leads to unprecedented pressure on coral reefs. Understanding the effects and implications of coral bleaching, and identifying strategies to reduce stress and mitigate impacts, are urgent challenges for the conservation and management of coral reefs worldwide. The Great Barrier Reef Marine Park (GBRMP) has experienced two major coral bleaching events in recent years: 1998 and 2002. The spatial extent of these events, combined with the high level of mortality seen at the most severely affected sites, has lead to widespread concern about the future of the Great Barrier Reef in the face of global climate change. Marshall, P and Schuttenberg, H. (2006) A reef manager’s guide to coral bleaching. Great Barrier Reef Marine Park Authority: Townsville (Australia), 163 pp. The biological diversity and productivity of coral reefs underpins the welfare of many societies throughout the world's tropical regions. Coral reefs form the foundation of dive tourism industries, support fisheries and are important to cultural traditions. They play an essential role in buffering coastal communities from storm waves and erosion, and they contain a largely untapped wealth of biochemical resources. Tens of millions of people depend upon reefs for all or part of their livelihood, and over a billion people rely on reef-related fisheries worldwide. With coral reefs providing such essential services to humans, the prospect of their continued widespread degradation is of concern. Pollution, habitat destruction, disease and unsustainable fishing have now led to declines in reef condition throughout the world. Against this backdrop of conventional stresses, the threat of mass coral bleaching has recently emerged, leading to what has been widely acknowledged as a 'coral reef crisis'. Mass coral bleaching has affected hundreds to thousands of kilometres of reefs simultaneously. It has caused stress, and in many cases extensive coral mortality, to nearly every coral reef region. In 1997-98 alone, mass bleaching is estimated to have caused over 90 per cent coral mortality to 16 per cent of the world's coral reefs. While strong initial signs of recovery have been observed in some locations, many will take decades to fully recover. Scientists agree that tropical seas will continue to warm over coming decades, increasing both the probability and severity of mass bleaching events. These scenarios pose particular challenges to coral reef managers, not the least because the main cause of mass coral bleaching–anomalously warm sea temperatures–is largely beyond their control. Yet, managers can play a critical role in helping reefs survive the threat of coral bleaching. Managers are in a unique position to increase our understanding of the phenomenon of coral bleaching, to take meaningful action during a bleaching event, and to develop strategies to support the natural resilience of reefs in the face of long-term changes in climate. Because of increasingly strong collaborations between reef managers and scientists, strategies

are being developed to directly address the threat of coral bleaching. Management needs and preliminary strategies were first documented in 2000, when the IUCN published Management of Bleached and Severely Damaged Coral Reefs. In 2002, the US Coral Reef Task Force called for a collaborative effort to identify actions local managers could take to address the impacts of climate change and mass bleaching on coral reefs. In response, three US government agencies (the National Oceanic and Atmospheric Administration, Environmental Protection Agency, and the Department of the Interior) convened an international workshop entitled 'Coral Reefs, Climate Change and Coral Bleaching' in June 2003. This workshop significantly advanced thinking about the strategies that could support managers in their efforts to respond to coral bleaching. Around the globe efforts have now begun to improve the prospects of coral reefs by: (1) identifying resilient areas and enhancing their protection, such as in Palau; and (2) implementing strategies to support ecosystem resilience. The Australian Government has implemented strategies to improve the protection of Australia’s Great Barrier Reef, including an integrated catchment management scheme and a new comprehensive zoning plan that protects unique areas and biodiversity, and includes 33% of the Great Barrier Reef in no-take areas. Further initiatives to build resilience principles into practical management of coral reefs and other marine ecosystems are under way, including efforts in Florida, USA. This guide builds on these recent initiatives by bringing together the latest scientific knowledge and management experience to assist managers in responding effectively to mass coral bleaching events. It synthesises science and management information, explores emerging strategies, and informs the ways managers deal with the complex human dimensions of these issues. Importantly, this guide is designed to provide pragmatic, science-based suggestions for adaptive management in this time of change.We commend it to reef managers worldwide, and hope that the experience of implementing the ideas within will further advance scientific knowledge and the practice of coral reef management. Mascia, MB. (2001) Designing effective coral reef marine protected areas. A synthesis report based on presentations at the 9th International Coral Reef Symposium: Bali (Indonesia), 22 pp. Coral reef ecosystems provide direct and indirect benefits to millions of people around the world. The long-term sustainability of these benefits is threatened, however, by human activities that impact reefs and reef ecosystems. Traditional efforts to manage human activities and protect coral reefs have proven inadequate, spurring calls for a more ecosystem-oriented approach. Central to this ecosystem-oriented approach to coral reef management is the establishment of marine protected areas (MPAs), a family of spatially-explicit marine management systems that includes underwater parks, fishery reserves, and wildlife sanctuaries. The promise of MPAs as a management tool has yet to be fully realized, in part because the science underlying effective MPA development and management is poorly understood. At the 9th International Coral Reef Symposium (ICRS) in Bali, Indonesia, dozens of scientists and practitioners presented cutting edge research on coral reef MPAs. This report synthesizes the findings from seventy-four ICRS presentations on MPAs and identifies select MPA policy and management implications from this natural and social scientific research.

Presentations at the ICRS underscored the scientific uncertainty that surrounds the biophysical design of MPAs, but provided some basic "rules of thumb" for MPA policymakers. There was general consensus that MPAs should be designated in high quality habitats, either in the midst of ocean gyres or in 'upstream' locations. Results gave little substantive guidance regarding the proper size for a functional MPA, though some interesting hypotheses did emerge. Researchers indicated that MPAs are more likely to function as relatively independent units than interdependent ecological systems, especially over large spatial scales. Biological performance was not correlated with the spatial extent of coral reef MPAs, suggesting that bigger is not necessarily better. ICRS presentations provided valuable insights into the sociopolitical characteristics of effective coral reef MPAs. Presenters stressed that MPAs are not a panacea, but rather dependent upon the larger matrix of coral reef management initiatives. If adjacent areas are not well managed, MPAs will likely be insufficient to maintain productive coral reef ecosystems. Presenters demonstrated that devolving authority for MPA development and management to local governments, user groups, and nongovernmental organizations spurs MPA establishment and enhances MPA management effectiveness. Collaborative MPA management structures, however, appear to offer the greatest potential for linking national resources with local interests and knowledge. Presentations regarding MPA regulatory systems identified emerging "best practices". One of the most contentious debates at the ICRS was whether MPAs should be "no-take" or permit limited extractive use. Though there was no resolution on this point (the answer seems to be "it depends on the situation"), presenters did agree that the rules governing resource use within coral reef MPAs must be clear, easily understood, and easily enforceable. Likewise, internal and external MPA boundaries must be easily recognized by resource users and by enforcement personnel. Presenters generally agreed that MPA decisionmaking must be an adaptive and broadly participatory process. Such processes permit social learning, build on diverse sources of knowledge, build trust, and enhance the legitimacy of MPA rules and regulations. Exactly how and when participation should occur was a matter of contention. MPA advisory committees were viewed as one appropriate mechanism for ongoing stakeholder participation in MPA development and management. Presenters emphasized that mechanisms must be established to ensure that stakeholder representatives are accountable and responsive to their constituents. Finally, presenters noted that differences among stakeholders with respect to their beliefs (i.e., perceptions of how the world works), values (i.e., perceptions of what is good, desirable, or just), and interests (i.e., desired outcomes) often hinder MPA development and management, reflecting the need for decision-makers to agree on process before trying to decide outcomes. Discussion of the management and administrative dimensions of MPAs was limited at the ICRS. Presenters noted that devolution of authority for enforcement could enhance capacity, and stressed the need to design enforcement systems that promote accountability among enforcers and appropriate (not draconian) penalties for noncompliance with MPA rules and regulations. Presenters suggested that clear management goals and objectives, as well as environmental education and outreach initiatives, facilitate effective MPA management. Research and monitoring were seen as critical components of MPA management, and speakers stressed the importance of monitoring both biological and social performance indicators. Speakers also stressed the importance of collecting baseline data, and sampling at multiple spatial

and temporal scales, in order to inform site development, measure change over time, and provide the basis for adaptive management. Finally, speakers noted that enlisting stakeholders in the collection and analysis of research and monitoring data educates participants and builds capacity and trust. During the ICRS, special emphasis was placed upon the role of no-take MPAs in supporting sustainable coral reef fisheries in Southeast Asia. Community participation, sustainable financing, enforcement, planning and design, and adaptive management were identified as five critical challenges to the development and management of MPAs in the region. At an evening workshop sponsored by NOAA, IUCN, and The Nature Conservancy, participants identified priority actions that would enhance MPA management across the region. These priority actions included training in community-based management, a regional inventory of experiences with sustainable financing, the development of model legislation and policy frameworks for decentralized enforcement, a regional assessment of priority sites for no-take MPAs, and the development of adaptive management pilot projects. Readers of this report should recognize its limitations. The scientific synthesis presented herein is based on the notes of nine volunteers and the author, who used a standard form to characterize seventy-four ICRS presentations most relevant to MPA development and management. The author synthesized these notes into the report summary, and derived policy implications from the report summaries and his personal knowledge of the natural and social scientific literature. These methods may have introduced uncertainties or biases into the report. Furthermore, neither the research upon which this report is based nor this report itself has been peer-reviewed, and therefore this report does not merit the same level of confidence as refereed research. McClanahan, TR. (2000) Bleaching damage and recovery potential of Maldivian coral reefs. Marine Pollution Bulletin 40, 587-597. Same-site comparisons of Maldive-Chagos reefs in the 1990s with studies before 1980 have found large losses in coral cover that were probably associated with warm El Niño events. To determine the spatial extent of this damage and potential for recovery I surveyed benthic cover, hard coral communities, and coral recruitment in previously unsurveyed sites in three reef atolls of the central Maldives in 1999, one year after the warmest recorded El Niño event of 1997-1998. Coral cover was the lowest recorded for this region, at 8%, and evidence for the local extirpation of species was found. Most reefs are presently dominated by coralline and turf algae (68%) with erect fleshy algae and sponge being higher than previously reported on Maldivian reefs. Branching coral species appeared most affected and the dominant coral genera in 1998 were massive Porites and Astreopora, whereas the original community was dominated by Acropora. Coral recruits were sufficiently abundant, at 29 ind/m2, to insure recovery of coral cover, but the most common recruits were in genera previously reported as subordinate genera, such as Pavona (11.7 ind/m2) and Coscinarea (4.4 ind/m2), whereas the previously dominant branching and encrusting species (Acropora, Montipora and Pocillopora) had recruit densities less than 0.65 ind/m2. Unless there is significant compensation in growth and mortality there may be future changes in coral species composition and benthic cover of these reefs.

McClanahan, TR, Muthiga, NA and Mangi, S. (2001) Coral and algal interactions after the 1998 coral bleaching: Interaction with reef management and herbivores in Kenyan reefs. Coral Reefs 19, 380-391. Interaction between the El Niño and Indian Ocean dipole ocean atmosphere quasi-periodic oscillations produced one of the warmest seawater temperatures on record in 1998. During the warm northeast monsoon in March and April, Kenya's shallow coral reefs experienced water temperatures between 30 and 31°C and low winds. This caused large-scale bleaching of hard and soft corals at the end of March, which extended into the cooler months of May and June. Direct observations of coloration in the Mombasa Marine National Park found that the coral genera Acropora, Millepora, Pocillopora, branching Porites and Stylophora showed rapid bleaching and high mortality by the end of May 1998. Other hard coral genera that bleached significantly included Echinopora, Favia. Favites, Galaxea, Hydnophora, Goniopora, Leptoria, Montipora, Platygyra and massive Porites, but mortality was variable among these genera. Astreopora, Coscinarea, Cyphastrea and Pavona were the least responsive genera, with some paling, but little evidence of full bleaching or significant mortality. We compared changes in reef ecology in four national parks (protected from fishing) with four non-park areas (heavy fishing) to determine how coral mortality and herbivory interact under the two management regimes. Benthic studies using line transects in 16 sites spread across ~150 km of coastline were completed before and 6 to 13 months after the bleaching event and found that the cover of nine hard coral genera including Acropora, Alveopora, Favites, Goniopora, Platygyra, Pocillopora, branching Porites, Stylophora and Tubipora decreased significantly (p<0.04) after the event, usually by >85%, and soft coral cover decreased by ~75%. One year after the bleaching, sites in the national parks experienced 88 and 115% increases in turf and fleshy algal cover, respectively, while reefs outside the parks had a 220% increase in fleshy algal cover with no appreciable change in turf-forming algal cover. There was, however, high spatial variation and no statistically significant difference in the change in fleshy algal cover between sites in and out of the national parks. Estimates of herbivory by both fish and sea urchins at each site were a good predictor of the change in fleshy algal cover over the 1-year post-bleaching period in about half of the sites. The other half of the sites did not exhibit large changes in fleshy algal cover, and algal cover at these sites was not clearly influenced by herbivory. The number of coral genera per transect at each site decreased significantly by 31 and 44% in and out of the national parks respectively. Larger-scale search sampling to determine the presence/absence of genera in study sites found consistent losses of coral genera from sites, but produced smaller differences than the line-transect method, suggesting that some genera persisted, but at very low population densities and small colony sizes. McClanahan, TR. (2002) The near future of coral reefs. Environmental Conservation 29, 460-483. In this paper the current status of coral reefs, predictions concerning the ecological state of coral reefs to the 2025 time horizon and the research needs that can help understanding and management activities that might alleviate detrimental ecological changes are evaluated and discussed. The present rate of CO2 emissions will produce an atmospheric concentration in 100 years not experienced during the past 20 million years and water temperatures above those of the past interglacial 130,000

years before present. Human influences on water temperatures, seawater chemistry (toxic substances, nutrients and aragonite saturation), the spread of diseases, removal of species and food web alterations are presently changing reef ecology. A significant ecological reorganization is underway and changes include a reduction in calcifying and zooxanthellae-hosting organisms, their obligate symbionts, and species at higher trophic levels, with an increase in generalist species of low trophic level that are adapted to variable environments. Late-successional fleshy brown algae of low net productivity or non-commercial invertebrates such as sea urchins, starfish and coral-eating snails will dominate many reefs. These changes will be associated with a loss of both net benthic and fisheries production and inorganic carbonate deposition; this will reduce reef complexity, species richness, reef growth and increase shoreline erosion. To avert these changes management is needed at both global and local levels. Both levels need to reduce greenhouse gases and other waste emissions and renew efforts to improve resource management including restrictions on the use of resources and globalization of resource trade, run-off and waste production, and balancing potential reef production and resource consumption. McClanahan, TR, Polunin, N and Done, T. (2002) Ecological states and resilience of coral reefs. Conservation Ecology 6. We review the evidence for multiple ecological states and the factors that create ecological resilience in coral reef ecosystems. There are natural differences among benthic communities along gradients of water temperature, light, nutrients, and organic matter associated with upwelling-downwelling and onshore-offshore systems. Along gradients from oligotrophy to eutrophy, plant-animal symbioses tend to decrease, and the abundance of algae and heterotrophic suspension feeders and the ratio of organic to inorganic carbon production tend to increase. Human influences such as fishing, increased organic matter and nutrients, sediments, warm water, and transportation of xenobiotics and diseases are common causes of a large number of recently reported ecological shifts. It is often the interaction of persistent and multiple synergistic disturbances that causes permanent ecological transitions, rather than the succession of individual short-term disturbances. For example, fishing can remove top-level predators, resulting in the ecological release of prey such as sea urchins and coral-eating invertebrates. When sea urchins are not common because of unsuitable habitat, recruitment limitations, and diseases, and when overfishing removes herbivorous fish, frondose brown algae can dominate. Terrigenous sediments carried onto reefs as a result of increased soil erosion largely promote the dominance of turf or articulated green algae. Elevated nutrients and organic matter can increase internal eroders of reef substratum and a mixture of filamentous algae. Local conservation actions that attempt to reduce fishing and terrestrial influences promote the high production of inorganic carbon that is necessary for reef growth. However, global climate change threatens to undermine such actions because of increased bleaching and mortality caused by warm-water anomalies, weakened coral skeletons caused by reduced aragonite availability in reef waters, and increased incidence of diseases in coral reef species. Consequently, many coral reefs, including those that are heavily managed, have experienced net losses in accumulated inorganic carbon in recent decades and appear likely to continue this trend in coming decades. Reefs urgently need to be managed with a view to strengthening their resilience to the increased frequency and intensity of these pressures. Ecological targets must include the restoration or maintenance of species

diversity, keystone species, spatial heterogeneity, refugia, and connectivity. Achieving these goals will require unprecedented cooperative synergy between human organizations at all political levels, from intergovernmental to local. McClanahan, TR and Maina, J. (2003) Response of coral assemblages to the interaction between natural temperature variation and rare warm-water events. Ecosystems 6, 551-563. We examined changes in coral assemblages in four back-reef locations across the warm 1998 E1 Niño–Southern Oscillation (ENSO) event based on annually collected line-transect data from 3 years before and after this event. The physical locations of the reefs differed such that there was a 120%–275% warm-season range in the SDs of seawater temperatures but only minor differences in mean temperatures, based on 2 non-ENSO years. We tested the predictions that (a) rare warm-water events would produce fewer changes in eurythermal than stenothermal coral assemblages; and (b) after the disturbance, the stenothermal assemblages would more closely resemble the eurythermal ones. The 1998 event produced fewer changes in coral cover and community similarity among the assemblages in the reefs with high variation in temperature than in those with low variation in temperature. Despite the initially lower taxonomic richness in the eurythermal assemblage, there was an additional loss of taxonomic richness in the high and none in the stenothermal reefs. There was some evidence for taxonomic convergence, of the stenothermal towards the eurythermal reefs and a general loss of some of the branching taxa, such as branching Porites, Pavona, and Stylophora, and a relative increase in massive Porites and Favia. There was, however, moderate site specificity that did not produce true convergence. The eurythermal assemblages maintained the basic community structure but lost taxonomic richness, whereas the opposite was true for the stenothermal assemblages. McClanahan, TR. (2004) The relationship between bleaching and mortality of common corals. Marine Biology 144, 1239-1245. Reef corals are likely to have many subtle but four gross responses to anomalous warm water. These are (1) not bleach and live (mortality <10%), (2) not bleach and die (mortality >20%), (3) bleach and live, and (4) bleach and die. The frequency of these four possible gross responses was determined for 18 common coral taxa over an exceptionally warm 1998 El Niño where intense bleaching was observed, and mortality determined from line transects averaged 41.2±34.7 (±SD). Field studies included (1) recording the loss of color (bleaching) and observing recently dead individuals among 6,803 colonies during five sampling periods and (2) estimating mortality based on 180 m of line-intercept transects completed 4 months before and near the end of the bleaching episode. There was no clear relationship between the loss of color and either direct observation or transect-based estimates of mortality for the 18 taxa. The morphology of the taxa did not influence color loss but branching and encrusting taxa had higher mortality than massive and submassive taxa. Loss of color and mortality are the most common responses to warm water as only Pavona did not lose color or die and only two taxa, Cyphastrea and Millepora, did not significantly lose color but died. Of the 15 taxa that lost color, five taxa, Astreopora, Favia, Favites, Goniopora, and Leptoria, did not die. These taxa are those most likely to have reduced potential mortality by the loss of pigments and associated algal

symbionts. Death of the branching taxa was detected reasonably by direct field observation but some taxa were underestimated when compared with mortality estimates based on line transects. Death of encrusting and massive taxa including Echinopora, Galaxea, Hydnophora, Montipora, Platygyra, and massive Porites was poorly detected from direct observations but they proved to have modest to high mortality (20–80%) based on line transects. There was no single response of these common corals to warm water but these data, collected during an extreme warmwater anomaly, indicate that the loss of color is most frequently a sign of morbidity, particularly for branching and encrusting taxa. McClanahan, TR, Baird, AH, Marshall, PA and Toscano, MA. (2004) Comparing bleaching and mortality responses of hard corals between southern Kenya and the Great Barrier Reef, Australia. Marine Pollution Bulletin 48, 327-335. We compared the bleaching and mortality response (BMI) of 19 common scleractinian corals to an anomalous warm-water event in 1998 to determine the degree of variation between depths, sites, and regions. Mombasa corals experienced a greater temperature anomaly than those on the Great Barrier Reef (GBR) sites and this was reflected in the greater BMI response of most taxa. Comparing coral taxa in different sites at the same depth produced high correlation coefficients in the bleaching response in Kenya at 2 m (r = 0.86) and GBR at 6 m depth sites (r = 0.80) but less in the GBR for shallow 2 m sites (r = 0.49). The pattern of taxa susceptibility was remarkably consistent between the regions. Coral taxa explained 52% of the variation in the response of colonies to bleaching between these two regions (Kenya BMI = 0.90 GBR BMI +26; F(1,9) = 18.3; p < 0:001; r2 = 0.52). Stylophora and Pocillopora were consistently susceptible while Cyphastrea, Goniopora Galaxea and Pavona were resistant in both regions. Three taxa behaved differently between the two regions; Acropora, and branching Porites were both moderately affected on the GBR but were highly affected in Kenya while the opposite was true for Pavona. These results suggest that a colonies response to bleaching is phylogenetically constrained, emphasizing the importance of features of the host’s physiology or morphology in determining the response to thermal stress. McClanahan, TR, Maina, J, Moothien-Pillay, R and Baker, AC. (2005) Effects of geography, taxa, water flow, and temperature variation on coral bleaching intensity in Mauritius. Marine Ecology Progress Series 298, 131-142. This study describes the response of common coral taxa at 15 sites to a warm-water anomaly in Mauritius in March 2004. Sites circumscribed the island and differed in their water flow and thermal history as a result of variation in local current patterns. We observed 2 distinct positive responses of coral taxa to the anomaly that correlated with their local abundance, with a group of sparse (22 taxa covering <5 cm m–1) and abundant taxa (7 taxa covering >5 cm m–1). The 2 most dominant taxa Acropora and Montipora were among the most susceptible genera in the abundant group, while Seriatopora and Alveopora were the most susceptible taxa in the sparse group. This suggests that a temperature anomaly that is sufficient to cause mortality will remove taxa from 2 positions in the community spectrum with consequences for both ecological functions and diversity. We found that bleaching intensity at the sites was positively associated with water flow, with the most intense bleaching and highest currents on the windward and offshore sites. The algal symbiont communities

in nearly all of the corals sampled on both sides of the island and 2 depths were dominated by diverse Symbiodinium in Clade C, indicating that the observed differences in response among coral taxa and sites were unlikely to be greatly affected by the types of symbionts they contained. We suggest that high water flow reduces background stress and acclimation, and results in corals that are less tolerant of rare temperature anomalies. McClanahan, TR, Maina, J, Starger, CJ, Herron-Perez, P and Dusek, E. (2005) Detriments to post-bleaching recovery of corals. Coral Reefs 24, 230-246. Predicting the response of coral reefs to largescale mortality induced by climate change will depend greatly on the factors that influence recovery after bleaching events. We experimentally transplanted hard corals from a shallow reef with highly variable seawater temperature (23–36°C) to three unfished marine parks and three fished reefs with variable coral predator abundance and benthic cover. The transplanted corals were fragmented colonies collected from a reef that was relatively undisturbed by the 1997–1998 warm-water temperature anomaly, one of the most extreme thermal events of the past century, and it was assumed that they would represent corals likely to succeed in the future temperature environment. We examined the effects of four taxa, two fragment sizes, an acclimation period, benthic cover components, predators and tourists on the survival of the coral fragments. We found the lowest survival of transplants occurred in the unfished marine parks and this could be attributed to predation and not tourist damage. The density of small coral recruits approximately 6 months after the spawning season was generally moderate (~40–60/m2), and not different on fished and unfished reefs. Coral recovery between 1998 and 2002 was variable (0–25%), low (mean of 6.5%), and not different between fished and unfished reefs. There was high variability in coral mortality among the three unfished areas despite low variation in estimates of predator biomass, with the highest predation occurring in the Malindi MNP, a site with high coralline algal cover. Stepwise multiple regression analysis with 14 variables of coral predators and substratum showed that coralline algae was positively, and turf algae negatively associated with mortality of the transplants, with all other variables being statistically insignificant. This suggests that alternate food resources and predator choices are more important than predator biomass in determining coral survival. Nonetheless, large predatory fish in areas dominated by coralline algae may considerably retard recovery of eurythermal corals. This will not necessarily retard total hard coral recovery, as other more predator-tolerant taxa can recover. Based on the results, global climate change will not necessarily favor eurythermal over stenothermal coral taxa in remote or unfished reefs, where predation is a major cause of coral mortality. McClanahan, TR, Verheij, E and Maina, J. (2005) Comparing the management effectiveness of a marine park and a multiple-use collaborative fisheries management are in East Africa. Aquatic Conservation: Marine and Freshwater Ecosystems. 1. The coral reefs across the international border between Kenya and Tanzania, where historical differences in government policy and socio-economic conditions created two different management systems, were examined: a large permanent closed area and a collaborative fisheries management project that used gear

management and small voluntarily and temporary closed areas, respectively. The diversity and ecology of the reefs in these two management systems were compared spanning a seven-year period to evaluate the effectiveness of the management and to assess the ecological response to a large-scale water-temperature anomaly in 1998. 2. Comparisons of rates of predation on sea urchins and of herbivory, using a seagrass assay, were made along with measures of benthic cover and fish abundance and diversity. 3. The collaborative fisheries management system was successful in increasing fish stocks, reducing erect algae, and maintaining ecological diversity and stability across the thermal anomaly. This management system, however, was not successful in protecting the expected full biodiversity of fish, predation rates on sea urchins, or the sensitive, branching coral species. Management of the fishery also increased fish stocks in the adjacent, large, permanently closed area, compared to Kenyan parks without this management. 4. The large, permanently closed area in the other system maintained high diversity, high predation rates on sea urchins and high herbivory rates, which maintained erect algae abundance and diversity at low levels. The temperature anomaly was destructive to a number of the dominant delicate branching coral species, but overall coral cover and diversity were maintained, although dominance switched from branching Porites spp. to Seriatopora spp. over this period. The large closed area system protected the undisturbed ecology of these reefs and associated ecological processes, and the full diversity of fish and coral, including sensitive species such as branching corals and slow-growing fish. 5. Collaborative fisheries and large permanent closed area management have different attributes that, when combined, should achieve the multiple purposes of sustainable fisheries, ecosystem functions and protection of fishing-sensitive species. McClanahan, TR, Ateweberhan, M, Ruiz Sebastián, C, Graham, NAJ, Wilson, SK, Bruggemann, JH and Guillaume, MMM. (2007) Predictability of coral bleaching from synoptic satellite and in situ temperature observations. Coral reefs. Satellite and compiled in situ observations of sea surface temperatures have greatly increased the ability to detect anomalous and persistent warm water and are being widely used to predict climate change, coral bleaching and mortality. A field-based synoptic view of coral bleaching spanning eight countries and ~35° of latitude in the western Indian Ocean tested the accuracy of synoptic temperature data derived from satellites and shipboard data to detect and predict bleaching during 2005. The ability to predict the degree of bleaching based on degree heating weeks data was moderate, but increased when past temperature anomalies and coral community susceptibility were included. It is estimated that slightly more than half of the bleaching response is due to anomalous warm water and nearly half due to taxa and community level acclimation or adaptation, where these two factors have opposing effects. Cumulative temperature anomalies do identify general areas with bleaching but both large over and underestimates of bleaching intensity were observed. Consequently, field observations are needed to confirm the synoptic satellite predictions for particular reefs, particularly where acclimation and reorganization of the coral community have occurred due to past bleaching events.

McCook, LJ, Jompa, J and Diaz-Pulido, G. (2001) Competition between corals and algae on coral reefs: A review of evidence and mechanisms. Coral Reefs 19, 400-417. Despite widespread acceptance that competition between scleractinian corals and benthic algae is important to the structure of coral reef communities, there is little direct experimental evidence that corals and algae do compete, and very little data on the processes and causality of their interactions. Most available evidence is observational or correlative, with intrinsic risks of confounded causality. This paper reviews and categorises the available evidence, concluding that competition between corals and algae probably is widespread on coral reefs, but also that the interaction varies considerably. Widespread replacement of corals by algae may often indicate coral mortality due to external disturbances, rather than competitive overgrowth, but may lead to competitive inhibition of coral recruitment, with consequences for reef recovery. We list eight specific processes by which corals and algae may affect each other, and suggest life history properties that will influence which of these interactions are possible We propose a matrix for algal effects on corals, which lists the subset of processes possible for each combination of coral life form and algal functional group. This table provides a preliminary framework for improved understanding and interpretation of coral-algal interactions. McLeod, E and Salm, RV. (2006) Managing mangroves for resilience to climate change. IUCN and The Nature Conservancy: Gland (Switzerland), 63 pp. Global climate change is one of the greatest challenges that humans will face in this century. Although geological records show climatic changes throughout history, the present rate of global warming threatens the survival of entire ecosystems. Among the most at-risk ecosystems are mangroves, which are especially vulnerable to sea-level rise, but the good news is that not all coastlines with mangrove forests are projected to experience a rise in relative sea level. At sites that are projected to experience rising seas, mangrove ecosystems on low relief islands and those deprived of sediment are especially vulnerable. In contrast, mangroves ecosystems with ample sediment supplies and/or room to move inland are likely to survive projected rates of sea-level rise. Mangrove species have demonstrated different tolerances to changes in sea level, salinity, and storms. By understanding which mangrove stands are able to survive sea-level rise and other changes, natural resource managers can identify and protect refuges that self-seed and act as sources for seeding of future mangrove communities. This paper is an attempt to provide some considerations for conservation practitioners as they design conservation strategies for mangroves. These ideas build upon the concept of resilience that was developed by West and Salm (2003) to address coral bleaching. Resilience is the ability of a system to undergo, absorb, and respond to change and disturbance, while maintaining its functions (Carpenter et al. 2001). West and Salm (2003) outline several strategies to help managers identify: 1) reef areas that are naturally resistant to coral bleaching (i.e., resistant areas); and 2) reef areas where environmental conditions are likely to promote maximum recovery after bleaching mortality has occurred (i.e., resilient areas). West and Salm (2003) recommend that these key areas, where environmental conditions appear to boost resistance and resilience during and after large-scale bleaching events, be incorporated into networks of marine protected areas. Although these principles were

developed to address coral reefs and increases in sea temperature, similar principles of resilience can be applied to mangroves and sea-level rise. Building resilience into mangrove conservation plans requires an understanding of how mangroves will respond to climate changes, what factors help them survive these changes, and, consequently, which mangroves are most likely to survive these changes. This paper provides an overview of mangrove ecosystems, discusses the benefits of mangroves to people, and the human and global threats that compromise mangrove ecosystems. This document describes the impacts of climate change on mangroves and outlines tools and strategies that enhance mangrove resilience. McManus, JW, Reyes Jr, RB and Nañola Jr, CL. (1997) Effects of Some Destructive Fishing Methods on Coral Cover and Potential Rates of Recovery. Environmental Management 21, 69-78. Effects of fishing with explosives (blastfishing) and sodium cyanide and of anchor damage on live coral were investigated on a heavily exploited fringing reef in Bolinao, Philippines from 1987 to 1990. A simple balance-sheet model indicated that approximately 1.4%/yr of the hermatypic coral cover may have been lost to blasting, 0.4%/yr to cyanide, and 0.03%/yr to coral-grabbing anchors, the potential coral recovery rate reduced by about one third from 3.8%/yr in the absence of disturbances to 2.4%/yr. These figures are subject to considerable uncertainty due to compounding of errors during computation. Reefs with patchy coral cover are more susceptible to damage from blastfishing because of targeting by fishers. Reefs with smaller corals may have greater resilience, because each unit of radial colony growth contributes a greater per cent increase in areal cover. Blastfishing in particular may reduce resilience to natural perturbations, leading to assemblages of small, sparse corals and reduced patchiness. McManus, JW, Meñez, LAB, Kesner-Reyes, KN, Vergara, SG and Ablan, MC. (2000) Coral reef fishing and coral-algal phase shifts: Implications for global reef status. ICES Journal of Marine Science 57, 572-578. Coral reef fisheries support tens of millions of people, mostly in developing countries. Fishing on reefs can be classified into three stages: manageable, ecosystem-overfished, and Malthusian-overfished. Fishing with blasting devices and poisons is often associated with the third stage. Reductions in herbivory caused by overfishing may enhance the likelihood of organic pollution causing a coral–algal phase shift following major disturbances. However, cage studies indicate that reduction in herbivory can lead to the proliferation of algae even in the absence of eutrophication. A major concern with the widespread coral bleaching associated with the 1997–1998 El Niño event is the likelihood that reefs already stressed by overfishing and organic pollution may not return to coral dominance after severe bleaching. Clues to levels of fishing and to the potential to recover from disturbances include changes in the de-vegetated ‘‘haloes’’ around coral stands on reef flats and the differing spectral signatures of live coral, recently dead coral, and coral covered with red encrusting, green filamentous, or brown frondose algae. These clues may facilitate broad area assessments of shallow reef areas via aircraft, space shuttles, or satellites.

McManus, JW and Polsenberg, JF. (2004) Coral-algal phase shifts on coral reefs: Ecological and environmental aspects. Progress in Oceanography 60, 263-279. This paper briefly reviews coral–algal phase shifts on coral reefs, with particular regard to summarizing the exogenous and endogenous factors in support of a proposed conceptual model, and to identifying critical information gaps. A phase shift occurs on a coral reef when the cover of a substrate by scleractinian corals is reduced in favor of macroalgal dominance, and resilience of the former condition is retarded because of ecological processes and/or environmental conditions. The change is often, but not always, associated with a perturbation such as coral bleaching, outbreaks of a coral-eating species, or storm damage. The new state is generally associated with some combination of reduced herbivory (from disease and/or fishing) and nutrient enrichment, although the relative importance of these factors is under debate and may vary among locations and even across single reefs. Disturbances that result in a state of generally low biotic three-dimensional structural complexity often precede a phase shift. Following such a disturbance, the system will pass to a state of higher biotic structural complexity, with either macroalgae or coral dominating. As the community progresses towards larger and more three-dimensionally complex corals or macroalgae, it exhibits greater resistance to shifting dominance from one state to the other. Studies of the phase-shift phenomena have been generally conducted at scales that are small relative to the sizes and inherent variability of whole coral reefs and systems of reefs. There is an urgent need for studies aimed at quantifying and simulating cause and effect aspects of the phase shift, including human–environment coupling, particularly in support of coral reef decision-making. McNeil, BI, Matear, RJ and Barnes, DJ. (2004) Coral reef calcification and climate change: The effect of global warming. Geophysical Research Letters 31. Coral reefs are constructed of calcium carbonate (CaCO3). Deposition of CaCO3 (calcification) by corals and other reef organisms is controlled by the saturation state of CaCO3 in seawater (W) and sea surface temperature (SST). Previous studies have neglected the effects of ocean warming in predicting future coral reef calcification rates. In this study we take into account both these effects by combining empirical relationships between coral calcification rate and W and SST with output from a climate model to predict changes in coral reef calcification rates. Our analysis suggests that annual average coral reef calcification rate will increase with future ocean warming and eventually exceed pre-industrial rates by about 35% by 2100. Our results suggest that present coral reef calcification rates are equivalent to levels in the late 19th century and does not support previous suggestions of large and potentially catastrophic decreases in the future.

Meehan, WJ and Ostrander, GK. (1997) Coral bleaching: A potential biomarker of environmental stress. Journal of Toxicology and Environmental Health 50, 529-552. Coral bleaching refers to the loss of symbiotic algae by host corals, or to the loss of pigmentation by the algae themselves, causing corals to appear white or "bleached." Some corals may regain algae or pigmentation and survive, but when bleaching is severe the host coral dies. Coral bleaching events have increased dramatically in the last two decades, and coral reefs throughout the world have been extensively degraded as a result. This article reviews coral bleaching for investigators working in the field of toxicology and environmental health, a group of scientists not normally exposed to this issue. Several environmental stressors have been correlated with bleaching, including fluctuations in sea surface temperatures and salinity, increased sedimentation, increased solar radiation, and contaminants such as oil and herbicides. Molecular mechanisms of bleaching are only beginning to be investigated and are thus far poorly understood. Toxicologists have the potential to make significant contributions toward understanding anthropogenic aspects of coral bleaching and elucidating molecular mechanisms of this important environmental problem. Meesters, EH, Nieuwland, G, Duineveld, CA, Kok, A and Bak, RPM. (2002) RNA/DNA rations of scleractinian corals suggest acclimatisation/adaptation in relation to light gradients and turbidity regimes. Marine Ecology Progress Series 227, 233-239. RNA/DNA ratios are used in many organisms as an indicator of growth, biomass or metabolic functioning. We hypothesised that hermatypic corals, as they depend for growth largely on energy transferred from their algal symbionts, may show an effect in the RNA/DNA ratio with factors influencing irradiance. This was tested along 2 environmental gradients: in relation to depth (decreasing irradiance) and in relation to environmental degradation (increased turbidity). RNA/DNA ratios were determined by HPLC in coral samples collected over depth gradients in 3 islands off the coast of north-west Java, Indonesia. The reefs range from shallow reefs (depth <3 m) in very turbid water, which are close to eroded shores, to intermediate reefs (depth <6 m) further from the shore with less turbid water, to offshore clear water control reefs (depth <20 m). RNA/DNA ratio was shown to be negatively correlated with depth in all but the most turbid conditions, suggesting it is an expression of metabolic acclimatisation of the coral host-symbiont community with decreasing irradiance (i.e. photoadaptation). Comparing the RNA/DNA ratio in clear and turbid waters, the decrease with depth tended to be steeper over a shorter depth range under turbid conditions, analogous with the more rapid extinction of light in these conditions. The relationship between the RNA/DNA ratio and light was consistently higher under turbid conditions (p = 0.0037), indicating that RNA/DNA ratio had shifted to a higher level, possibly indicating a genetic adaptation in the metabolic functioning of corals in the turbid environment. The positive relation between light and RNA/DNA ratio would suggest the opposite, i.e. lower ratio under turbid conditions. The RNA/DNA ratio may provide a relatively reliable method to determine the metabolic functioning (health) of individual coral colonies.

Mellin, C, Andréfouët, S and Ponton, D. (2007) Spatial predictability of juvenile fish species richness and abundance in a coral reef environment. Coral Reefs. Juvenile reef fish communities represent an essential component of coral reef ecosystems in the current focus of fish population dynamics and coral reef resilience. Juvenile fish survival depends on habitat characteristics and is, following settlement, the first determinant of the number of individuals within adult populations. The goal of this study was to provide methods for mapping juvenile fish species richness and abundance into spatial domains suitable for micro and meso-scale analysis and management decisions. Generalized Linear Models predicting juvenile fish species richness and abundance were developed according to spatial and temporal environmental variables measured from 10 m up to 10 km in the southwest lagoon of New Caledonia. The statistical model was further spatially generalized using a 1.5-m resolution, independently created, remotely sensed, habitat map. This procedure revealed that : (1) spatial factors at 10 to 100-m scale explained up to 71% of variability in juvenile species richness, (2) a small improvement (75%) was gained when a combination of environmental variables at different spatial and temporal scales was used and (3) the coupling of remotely sensed data, geographical information system tools and point-based ecological data showed that the highest species richness and abundance were predicted long a narrow margin overlapping the coral reef flat and adjacent seagrass beds. Spatially explicit models of species distribution may be relevant for the management of reef communities when strong relationships exist between faunistic and environmental variables and when models are built at appropriate scales. Mesoamerican Barrier Reef Systems Project. (2003) Training manual on design and development of management plans for marine protected areas. MBRS: Belize City (Belize), 66 pp. The constant degradation of marine and coastal ecosystems creates the urgent need to establish reserves and marine protected areas which allow the conservation of biodiversity as well as improve the living conditions of the human communities which depend directly on them. In order to guarantee conservation, sustainable use of the marine and coastal natural resources and community development, in addition to establishing marine protected areas and reserves, it is essential to have the appropriate tools, trained personnel, infrastructure and basic equipment. Management Plans are one of the fundamental tools that guide management and define the terms of the ecosystems, the human communities and the administrative components of the plan, required in accordance with the management category. For this reason, the Mesoamerican Barrier Reef System Project organized and coordinated a regional training course on the design and development of Management Plans for Marine Protected Areas. During this training, support material was provided based on existing experiences. This material was enriched by the input from delegates of the four countries in the MBRS region, which ultimately resulted in the creation of the current document in the form of a manual. The process involved the participation of an international expert, Alejandro Arrivillaga Cortéz, PhD, who prepared the initial training material, conducted the training,

collected the input from the participants and developed the final version of this manual, incorporating the comments of the Project Coordination Unit (PCU.) The training included case studies, formal presentations, field trips, individual country presentations, assigned reading and dynamic discussions. Likewise, the development of the manual was based on the Training for Trainers in the Management of Protected Areas Manual, developed by UNEP at its Regional Coordination Unit in Kingston, Jamaica. Existing Management Plans for the 15 priority MPA’s within the MBRS transboundary zones were reviewed and individual management experiences shared by the participants were incorporated, to create a manual with the particular characteristics of the four countries involved in the Project. With this manual, the MBRS Project reaffirms its commitment to generate support material for the conservation and sustainable use of marine and coastal resources, with the participation of the different sectors of civil society, governmental and non-governmental organizations, promoting participatory forums for decision-making. It is our intention that this tool be used in mass training courses and serve as a guide in the processes of revising, updating, re-orienting and preparing Management Plans for MPA’s in the various countries of the Mesoamerican region. In so doing, it will strengthen the efforts to conserve biological diversity as well as foster community development by increasing the diversity of actors and levels of participation in the management process. Micheli, F and Halpern, BS. (2005) Low functional redundancy in coastal marine assemblages. Ecology Letters 8, 391-400. The relationship between species and functional diversity remains poorly understood for nearly all ecosystem types, yet determining this relationship is critically important for developing both a mechanistic understanding of community assembly and appropriate expectations and approaches to protecting and restoring biological communities. Here we use two distinct data sets, one from kelp forests in the Channel Islands, California, and one from a global synthesis of marine reserves, to directly test how variation in species diversity translates into changes in functional diversity. We find strong positive relationships between species and functional diversity, and increased functional diversity of fish assemblages coinciding with recovery of species diversity in marine reserves, independent of the method used for classifying species in functional groups. These results indicate that low levels of redundancy in functional species traits exist across a suite of marine systems, and that fishing tends to remove whole functional groups from coastal marine ecosystems. Miclat, EFB. (2002) Selecting priority areas for conservation in the Sulu-Sulawesi marine ecoregion. Proceedings of the IUCN/WCPA Taipei Conference: Taipei (Taiwan), 297-303. Sulu-Sulawesi is a transborder marine ecoregion in the epicenter of global marine biodiverisity. Prevailing modes of resource uses gravely threaten this ecoregion. Current conservation approaches and initiatives are insufficient to address threats that operate over a large scale. Thus, innovations in conservation must be adopted to abate threats and maintain and/or restore biodiversity in an ecoregional scale. The WWF’s program on ecoregion conservation, which ultimate goal is to conserve an

ecologically representative complement of the ecoregion’s biodiversity and processes, is being developed. The planning phase includes the development of a methodology for identifying priorities for conservation in a large area of land or water. The use of the method in Sulu-Sulawesi resulted in the identification of 58 priority areas which when established into a network of protection will ensure the conservation of the fullest range possible of biodiversity in this globally significant marine ecoregion. Miller, K and Mundy, C. (2003) Rapid settlement in broadcast spawning corals: Implications for larval dispersal. Coral Reefs 22, 99-106. Broadcast spawning of gametes with planktonic development of larvae is the most common reproductive mode in tropical corals, and is generally thought to optimize the dispersal potential of larvae. To this end, many previous studies of coral larval dispersal have focused on the maximum time larvae can remain competent to settle and consequently how far they might disperse. However, dispersal ability of broadcastspawned coral larvae will be linked, at least in part, to the minimum time to settlement competency as well as the length of the planktonic period – although estimates of minimum time to competency remain largely anecdotal, with few rigorous studies of the pre-competent period. To determine the minimum time to larval settlement in two species of broadcast-spawning coral (Platygyra daedalea and Goniastrea favulus), we monitored larval settlement rates in aquaria every 6 h from the time larvae commenced swimming (i.e. were ciliated, fully developed larvae) for a period of approximately 10 days. For P. daedalea, peak settlement occurred between 60 and 66 h following fertilization (2.5 and 2.75 days), which is markedly earlier than the 4- to 6-day time period commonly cited as the minimum time before broadcast-spawned coral larvae are competent to settle. Surprisingly, it was also clear from our experimental results that settlement in P. daedalea occurred as a distinct pulse during the 60- to 66-h period, rather than continuously throughout the study period. G. favulus larvae also appear to be able to settle quickly (from 54 h following fertilization). We argue, on the basis of these short competency times and apparently rapid settlement, that dispersal in broadcast-spawning coral larvae may not be as great as has previously been assumed. Mitchelmore, CL, Schwarz, JA and Weis, VM. (2002) Development of symbiosis-specific genes as biomarkers for the early detection of cnidarian-algal symbiosis breakdown. Marine Environmental Research 54, 345-349. Coral bleaching, i.e. the loss of dinoflagellate symbionts from cnidarian hosts, is occurring globally at increasing rates, scales, and severity. The significance of these bleaching events to the health of coral reef ecosystems is extreme, as bleached corals exhibit high mortality, reduced fecundity and productivity and increased susceptibility to disease. This decreased coral fitness leads to reef degradation and ultimately to the breakdown of the coral reef ecosystem. To date there has been little work describing the application of biomarkers to assess coral health. The most commonly applied biomarker is, in fact, the bleaching event itself. We are interested in developing early warning biomarkers that can detect coral stress before bleaching occurs. Recently, several genes that are likely to function in regulating interactions between cnidarians and their symbionts have been characterized, using the temperate sea anemone Anthopleura elegantissima as a model species. One

"symbiosis gene" identified from the host genome, sym32, is expressed as a function of anemone symbiotic-state, where sym32 expression is higher in symbiotic cf. aposymbiotic (symbiont-free) anemones. Real-time quantitative RT-PCR suggested that the level of sym32 expression was correlated with the abundance of algae in the host. Furthermore, laboratory exposures of anemones to low levels of cadmium (0, 20, 100 microg(-1) CdCl2; 14 days), which caused no change in algal cell numbers, resulted in a down-regulation of sym32 compared to controls, indicating that sym32 expression may serve as a new sensitive early warning biomarker of cnidarian-algal symbiosis breakdown. Muko, S, Sakai, K and Iwasa, Y. (2001) Dynamics of marine sessile organisms with space-limited growth and recruitment: Applications to corals. Journal of Theoretical Biology 210, 67-80. We study a model for the community dynamics of marine sessile organisms with space limitation both in recruitment and in growth. We consider an open population in which recruits are supplied from a pelagic pool of larvae produced by adults in distant habitats. Assumptions are: the larval settlement rate is proportional to the amount of free space and to the abundance of larvae in the water column. The growth rate of settled individuals increases with the fraction of free space within the local habitat. We study the competition between two morphotypes with different rates of recruitment, growth, and mortality. When adult mortality is low, following a major disturbance that creates bare patch, the space is quickly filled by larval recruitment and adult growth. Then the morphotype composition changes slowly and converges to the equilibrium that is strongly affected by mortality. We also examine several other limiting cases in which one of the three demographic processes occurs either very slowly or very quickly. Based on the model behavior, we discuss the possible factors responsible for the spatial variation in the morphotype composition observed in coral communities. The dominance of branching corals in protected sites can be explained by their faster growth than tabular corals. The dominance of tabular corals in exposed sites can be explained either by lower mortality or by faster recruitment than branching corals. Muko, S, Sakai, K and Iwasa, Y. (2001) Size distribution dynamics for a marine sessile organism with space-limitation in growth and recruitment: Application to a coral population. Journal of Animal Ecology 70, 579-589. 1. Corals are clonal organisms and show a plastic growth. We study a partial differential equation model for the dynamics of size distribution of corals and predict the trajectory of recovery after a catastrophic disturbance, such as the recent bleaching that killed most corals in southern Japan. 2. We assume that the mean growth rate of colony size, measured in projected area, is a linear function of colony size, and that the variance in growth rate is proportional to the size, which is consistent with the growth data of a coral Acropora hyacinthus Dana 1846. 3. The model incorporates the space-limitation in colony growth and recruitment. The growth rate and recruitment rate are proportional to the fraction of free space within the local habitat. 4. In many corals, including A. hyacinthus, recruitment occurs in a short period once a year. However, our model illustrates that the colony size distribution does not show

distinguishable cohorts since the observed variance in growth rate is large. The model with discrete settlement and a large variance in growth rate results in size distributions that are very well approximated by an explicitly soluble model with constant recruitment and no growth variance. 5. When mortality is low, the dynamics of size distribution show two different phases in the recovery process. In the first phase, size distribution is determined by recruitment and growth, and can be predicted well by the case without mortality. After free space is depleted, recruitment and growth slow down and become balanced with mortality. The equilibrium size distribution is controlled by all the three processes. Both for the transient and the equilibrium size distribution, the average colony size increases with growth rate but decreases with recruitment rate. 6. Strongly skewed colony-size distributions in which small size classes have the largest numbers are generated for a wide range of parameters by the space-limitation in growth, even without partial death of colonies. Mumby, PJ, Chisholm, RM, Edwards, AJ, Andréfouët, S and Jaubert, J. (2001) Cloudy weather may have saved Society Island reef corals during the 1998 ENSO event. Marine Ecology Progress Series 222, 209-216. During the 1998 El Niño-Southern Oscillation (ENSO) event, mass coral bleaching in French Polynesia was patchy at a scale of 100s of km. Bleaching was extensive in parts of the Tuamotu archipelago (creating up to 99% coral mortality) but extremely mild in the Society Islands (Tahiti, Moorea), ca 350 km to the south-west, despite sea surface temperature (SST) anomalies being of similar magnitude to previous years in which mass bleaching occurred. We examine whether environmental variables account for this unexpected paucity of bleaching using a 50 yr record of SST, a 17 yr record of daily wind and cloud cover, and a 17 yr record of monthly sun hours. Records from Tahiti reveal that exceptionally high cloud cover significantly reduced the number of sun hours during the summer of 1998. Quadratic discriminant analyses of annual bleaching occurrence based on up to 3 predictors (cumulative degree heating months, wind speed, and cloud cover during periods of elevated summer SST) only predicted the correct bleaching scenario for 1998 when cloud cover was added to the function. The results demonstrate that the interactive effect of cloud cover can reverse the bleaching predictions of such statistical models. We suggest that reduced radiative stress, resulting from high cloud cover, may have prevented large-scale coral bleaching in 1998. Mumby, PJ, Chisholm, JRM, Edwards, AJ, Clark, CD, Roark, EB, Andréfouët, S and Jaubert, J. (2001) Unprecedented bleaching-induced mortality in Porites spp. At Rangiroa Atoll, French Polynesia. Marine Biology 139, 183-189. In April-May 1998, mass coral bleaching was observed in the lagoon of Rangiroa Atoll, Tuamotu Archipelago, French Polynesia. Six months later, the extent of bleaching-induced coral mortality was assessed at three sites. Corals in the fast-growing genus Pocillopora had experienced > 99% mortality. Many large colonies of the slow-growing genus Porites (mean horizontal cross-sectional area 5.8 m2) had also died - a phenomenon not previously observed in French Polynesia and virtually unprecedented world-wide, At one site, 25% of colonies, or 44% of the pre-bleaching cover of living Porites, experienced whole-colony mortality. At the two other sites, recently dead Porites accounted for 41% and 82% of the pre-bleaching live cover.

Mortality in Porites was negatively correlated with depth between 1.5 and 5 m. Using a 50-year dataset of mean monthly sea surface temperature (SST), derived from ship- and satellite-borne instruments, we show that bleaching occurred during a period of exceptionally high summer SST. 1998 was the first year in which mean monthly SSTs exceeded the 1961-1990 upper 95% confidence limit (29.4°C) for a period of three consecutive months. We suggest that the sustained 3-month anomaly in local summer SST was a major cause of coral mortality, but do not discount the synergistic effect of solar radiation. Recovery of the size-frequency distribution of Porites colonies to pre-bleaching levels may take at least 100 years. Mumby, PJ, Foster, NL and Glynn Fahy, EA. (2005) Patch dynamics of coral reef macroalgae under chronic and acute disturbance. Coral Reefs 24, 681-692. The patch dynamics (colonisation rate, growth rate, and extinction rate) are quantified for two dominant species of macroalgae on a Caribbean forereef in Belize: Lobophora variegata (Lamouroux) and Dictyota pulchella (Hörnig and Schnetter). Measurements were taken on time scales of days, weeks, months, and years during which three hurricanes occurred. All patches were followed on naturally occurring ramets of dead Montastraea annularis. The first hurricane (Mitch) caused massive coral mortality and liberated space for algal colonisation. The cover of Lobophora increased throughout the study and herbivores did not appear to limit its cover within a 4 year time frame. In contrast, the cover of D. pulchella fluctuated greatly and showed no net increase, despite an increase in parrotfish biomass and settlement space. Variation in the overall percent cover of an alga is not indicative of the underlying patch dynamics. The steady rise in the cover of Lobophora took place despite a high turnover of patches (12–60% of patches per year). The patch dynamics of Dictyota were slower (7–20%), but a greater patch density and threefold higher lateral growth rate led to greater fluctuations in total cover. The dynamics of algal patches are size-specific such that larger patches are less likely to become extinct during hurricanes.

Mumby, PJ. (2006) The impact of exploiting grazers (Scaridae) on the dynamics of Caribbean coral reefs. Ecological Applications 16, 747-769. Coral reefs provide a number of ecosystem services including coastal defense from storms, the generation of building materials, and fisheries. It is increasingly clear that the management of reef resources requires an ecosystem approach in which extractive activities are weighed against the needs of the ecosystem and its functions rather than solely those of the fishery. Here, I use a spatially explicit simulation model of a Caribbean coral reef to examine the ecosystem requirements for grazing which is primarily conducted by parrotfishes (Scaridae). The model allows the impact of fishing grazers to be assessed in the wider context of other ecosystem processes including coral-algal competition, hurricanes, and mass extinction of the herbivorous urchin Diadema antillarum. Using a new analytical model of scarid grazing, it is estimated that parrotfishes can only maintain between 10% and 30% of a structurally complex forereef in a grazed state. Predictions from this grazing model were then incorporated into a broader simulation model of the ecosystem. Simulations predict that scarid grazing is unable to maintain high levels of coral cover (> or = 30%) when severe hurricanes occur on a decadal basis, such as occurs in parts of the northern Caribbean. However, reefs can withstand such intense disturbance when grazing is

undertaken by both scarids and the urchin Diadema. Scarid grazing is predicted to allow recovery from hurricanes when their incidence falls to 20 years or less (e.g., most of Central and South America). Sensitivity analyses revealed that scarid grazing had the most acute impact on model behavior, and depletion led to the emergence of a stable, algal-dominated community state. Under conditions of heavy grazer depletion, coral cover was predicted to decline rapidly from an initial level of 30% to less than 1% within 40 years, even when hurricane frequency was low at 60 years. Depleted grazers caused a population bottleneck in juvenile corals in which algal overgrowth caused elevated levels of postsettlement mortality and resulted in a bimodal distribution of coral sizes. Several new hypotheses were generated including a region-wide change in the spatial heterogeneity of coral reefs following extinction of Diadema. The management of parrotfishes on Caribbean reefs is usually approached implicitly through no-take marine reserves. The model predicts that depletion of grazers in nonreserve areas can severely limit coral accretion. Other studies have shown that low coral accretion can reduce the structural complexity and therefore quality of the reef habitat for many organisms. A speculative yet rational inference from the model is that failure to manage scarid populations outside reserves will have a profoundly negative impact on the functioning of the reserve system and status of non-reserve reefs. Mumby, PJ, Hedley, JD, Zychaluk, K, Harborne, A and Blackwell, PG. (2006) Revisiting the catastrophic die-off of the urchin Diadema antillarum on Caribbean coral reefs: Fresh insights on resilience from a simulation model. Ecological Modelling 196, 131-148. In 1983, the dominant urchin of Caribbean coral reefs (Diadema antillarum) experienced massive disease-induced mortality and its functional extinction persists to this day. Concurrently, reports of coral disease, coral bleaching and nutrification began to appear and many Caribbean reefs have deteriorated in the last two decades. Here, we describe a spatial simulation model of physical and ecological processes occurring on a dominant Caribbean for reef habitat, often referred to as Montastraea reef. The model uses a square lattice to simulate the dynamics of two types of hard coral (a brooder and spawner), cropped algal substrata and macroalgae. Grazing occurs independently by Diadema and parrotfishes and influences the competitive interactions between corals and macroalgae. Four important types of disturbance are simulated: loss of the urchin Diadema, hurricanes, fishing of parrotfishes and nutrification. We use the model to revisit the impact of losing the keystone herbivore, Diadema. The urchin imparted great resilience to Caribbean reefs in the face of severe disturbances including: high hurricane frequencies (once per decade), overfishing of scarids and a range of nutrification levels, including a doubling of the vertical growth rate, vegetative growth rate and colonisation rate of macroalgae. Parrotfishes are able to compensate for the loss of this urchin under a limited range of physical and ecological conditions. When not exploited, parrotfishes allow coral to persist at least at equilibrial levels when hurricane frequencies drop to around once every 20 years (e.g. those found in Central America). However, coral cover declined under decadal hurricane frequencies which may partly explain observations in regions of intense hurricane activity such as Florida. Parrotfish grazing is sufficiently intense to mitigate nutrification impacts providing that the cover of corals is not heavily depleted by other processes such as hurricanes and disease. Indeed, the cover of living coral exhibits a

strong interaction with both grazing and nutrification in determining the survival of coral recruits. Reefs of higher coral cover have greater resilience for a given level of nutrification or grazing and therefore management efforts that arrest declines in coral cover will build greater natural resilience in the system. The model provides a strong case for conserving parrotfishes because reef decline is predicted to be inevitable when urchins are scarce and fishing of grazers is intense (e.g. to the levels seen in Jamaica). Lastly, the model provides a new perspective on the bottom-up versus top-down debate of macroalgal blooms in Jamaica. Nutrification impacts are predicted to be most important when urchins are scarce and parrotfishes are relatively lightly exploited. Therefore, a full understanding of the impact of nutrification will only occur if no-take marine reserves permit scarid biomass to reach unexploited levels. Under these conditions, coral cover is only predicted to increase if nutrification levels are low. Nadaoka, K, Nihei, Y, Wakaki, K, Kumano, R, Kakuma, S, Moromizato, S, Omija, T, Iwao, K, Shimoike, K, Taniguchi, H, Nakano, Y and Ikema, T. (2001) Regional variation of water temperature around Okinawa coasts and its relationship to offshore thermal environments and coral bleaching. Coral Reefs 20, 373-383. To clarify the relationship between the regional variability of water temperatures and the spatially non-uniform appearances of coral mass bleaching and consequent mortality, we have investigated the near-shore temperatures continuously monitored around the Ryukyu Islands in 1999 and NOAA-SST image data in 1998 and 1999. They indicate appreciable regional differences in temperature (e.g., at the Kerama Islands located southwest of Okinawa Island the temperature in summer was generally lower than that at the Okinawa Island coasts). Comparison of the solar radiation distribution and NOAA-SST images with bottom bathymetry and a numerical experiment suggest that the presence of the shelf around the Kerama Islands caused the relatively lower temperature (i.e., the warm water mass from the Kuroshio currents may be propagated toward Okinawa Island but may be blocked at the shelf edge, resulting in relatively lower temperature at the Kerama Islands). Further comparison with the reported changes in the coral coverage in the Ryukyu Islands after the mass coral bleaching event in summer 1998 suggests that one of the primary causes of the regional variability of the coral bleaching and consequent mortality is the regional variation of water temperature. Nakamura, T and van Woesik, R. (2001) Water-flow rates and passive diffusion partially explain differential survival of corals during the 1998 bleaching event. Marine Ecology Progress Series 301-304. In the western Pacific during 1998, coral bleaching, or the paling of corals through loss of pigmentation or loss of symbiotic algae (zooxanthellae), coincided with some of the warmest sea-surface temperatures (SSTs) on record. However, there was considerable spatial variation in coral survivorship; for example, corals of the same species at different locations around the Ryukyu Islands (Japan), within kilometers of each other, showed vastly different responses. Some locations experienced 100% coral mortality while other locations, nearby, suffered little coral mortality. Here we show experimental evidence for high survivorship of Acropora digitifera coral colonies that were subjected to both high SSTs (ranging from 26.22 to 33.65°C) and high-water flow (50 to 70 cm s–1), while corals that were subjected to both high SSTs and

low-water flow (2 to 3 cm s–1) showed low survivorship. All experiments were conducted under high irradiance (~95% photosynthetically active radiation). We also empirically show that no coral mortality occurred when SSTs were below 30°C (ranging from 26.64 to 29.74°C) under similar flow regimes. The spatial differences in coral mortality during the 1998 bleaching event may have been, in part, a result of differences in water-flow rates that induced differential rates of passive diffusion, which varied among habitats. Nakamura, T, Yamasaki, H and van Woesik, R. (2003) Water flow facilitates recovery from bleaching in the coral Stylophora pistillata. Marine Ecology Progress Series 256, 287-291. Exposing partially bleached samples of Stylophora pistillata coral colonies to controlled water-flow treatments revealed differential recovery rates that varied in accordance with flow rate. Recovery of the number of zooxanthellae per unit area and chlorophyll a concentrations increased rapidly in moderate-flow treatments (of 20 cm s–1), after an initial 3 wk of stasis. Colonies in low-flow treatments (≤3 cm s–1) remained pale for the entire 7 wk experimental period and showed only slight increases in zooxanthellae per unit area. In an earlier study, we showed that when corals were subjected to high temperature and high irradiance, coral bleaching occurred in low-flow but not in moderate-flow treatments; the present study shows that recovery from bleaching is facilitated by flow treatment. In combination, prevention of and rapid recovery from bleaching by enhanced water flow suggest that the mechanisms that lead to dysfunction of the algae-coral symbiosis are driven by mass-transfer-limited processes. Nakamura, T, van Woesik, R and Yamasaki, H. (2005) Photoinhibition of photosynthesis is reduced by water flow in the reef-building coral Acropora digitifera. Marine Ecology Progress Series 301, 109-118. While photosynthesis of symbiotic algae is essential for reef-building corals, excess irradiance inhibits photosynthesis through photoinhibition, which can lead to coral bleaching under elevated temperature conditions. Here we show that water flow reduces photoinhibition of in hospite endosymbionts in the coral Acropora digitifera. Diurnal monitoring of chlorophyll fluorescence, under 2 different flow regimes (<3 and 20 cm s–1 flow rates) in an outdoor aquarium, showed reduced photoinhibition, but only under moderate flow conditions (20 cm s–1). Experimental (laboratory) measurements, on time scales ranging from minutes to hours, showed that flow-mediated reductions in photoinhibition occurred not by enhancing recovery of the damaged photosystem, but rather through inducing differential photodamage. Moreover, experiments involving sequential light oscillations (500/20 and 1000/20 µmol photons m–2 s–1) at 3 flow regimes, <3, 10, and 20 cm s–1, on a time scale ranging from hours to days, revealed water-velocity-dependent reductions of dynamic photoinhibition. These results, on time scales ranging from minutes to weeks, confirm that reduced water flow amplifies photodamage of algal photosynthesis under strong irradiance, which in turn affects coral tolerance to strong irradiance and temperature.

Nellemann, C and Corcoran, E. (2006) Our precious coasts – Marine pollution, climate change and the resilience of coastal ecosystems. UNEP: Arendal (Norway), 38 pp. Climate change is seriously impacting the world’s marine ecosystems. Massive coral bleaching episodes have impacted the function of the reefs and increased rates of mortality. Coral reefs support over one million plant and animal species and their economic value is projected to more than US $ 30 billion annually. Extreme climatic conditions, however, are most likely to increase in the future with current climate scenarios. Projected increases in carbon dioxide and temperature exceed the conditions under which coral reefs have flourished over the past 500 000 years. Coral reefs are crucial biodiversity hotspots and support both coastal fisheries and tourism in many regions. Coral reefs, however, are in decline in many regions as a result of numerous pressures, including, but not limited to, extreme climate events, unsustainable fishing practices, diseases, sedimentation, and discharge of untreated sewage. Increasing resilience and securing rapid recovery of coral reefs will be essential for the ability of these ecosystems to support coastal fisheries and coastal livelihoods and cultures in the future. However, this resilience and recovery may be seriously impounded by unsustainable coastal infrastructure development and marine pollution from land-based sources. At the current rate of growth, coastal development may impact up to 90% of the tropical and temperate coastlines by 2032 if development continues unchecked. While progress has been made to reduce the discharge and impacts of oil spills and persistent organic pollutants (POP’s), there now needs to be a focus on the largest current threats to the coastal marine environment : untreated sewage and piecemeal coastal development. Over 90% of all the world’s coral reefs are found in the Indo-Pacific region of Asia, but also found here are some of the largest increases and levels of emissions of untreated sewage discharge and coastal marine pollution and development. A drastic increase in the appropriate integrated management of coastlines particularly near marine protected areas is urgently needed. Furthermore, an increase in enforcement and extent of protected coastlines, is urgently needed to secure the future diversity and recovery of coral reefs from climate change. Such combined joint protected areas may form source-“islands” or coral “treasure vaults” for re-colonization of damaged areas. Furthermore, the combined cumulative effects of coastal overfishing, marine pollution and coastal development may impact the long-term productivity of the coastal zone. This, in turn, may lower the capacity of these systems to support human livelihoods in the long-term, This challenge requires effective integrated landuse planning including fisheries, tourism and costal infrastructure development, as well as proper watershed management further inland. Ninio, R and Meekan, MG. (2003) Spatial patterns in benthic communities and the dynamics of a mosaic ecosystem on the Great Barrier Reef, Australia. Coral Reefs 21, 95-103. The benthic communities of the Great Barrier Reef (GBR) have been characterized as a mosaic with patches at scales of tens to hundreds of kilometers formed by clusters of reefs with comparable environmental settings and histories of disturbance. We use data sets of changes in cover of abundant benthic organisms to examine the relationship between community composition and the dynamics of this mosaic. Our data were compiled from seven annual video surveys of permanent transects on the

north-east flanks of up to 52 reefs at different shelf positions throughout most of the GBR. Classification analysis of these data sets identified three distinct groups of reefs, the first dominated by poritid hard corals and alcyoniid soft corals, the second by hard corals of the genus Acropora, and the third by xeniid soft corals. These groups underwent different amounts of change in cover during the period of our study. As acroporan corals are fast growing but susceptible to mortality due to predators and wave action, the group of reefs dominated by this genus displayed rapid rates of growth and loss of cover. In contrast, cover in the remaining groups changed very slowly or remained stable. Some evidence suggests that competition for space may limit growth of acroporan corals and thus rates of change in the group dominated by xeniid soft corals. These contrasting patterns imply that susceptibility to, and recovery from, disturbances such as cyclones, predators, and bleaching events will differ among these groups of reefs. Nishikawa, A and Sakai, K. (2005) Genetic connectivity of the scleractinian coral Goniastrea aspera around the Okinawa Islands. Coral Reefs 24, 318-323. Genetic variation and gene flow in the scleractinian coral Goniastrea aspera (Verrill), found around the north–south Okinawa Islands, were studied using allozyme and starch gel electrophoresis. The relative contribution of sexual and asexual reproduction to recruitment was determined. Analysis of multilocus genotypes of samples, collected at least 3 m apart, identified a high number of unique genotypes (NG) relative to the number of individuals sampled (N) (mean NG: N=0.97±0.03 [SD]), and also highly observed genotypic diversity (GO) relative to expected genotypic diversity under Hardy–Weinberg equilibrium (GE) (mean GO: GE=0.95±0.06 [SD]). These results suggest that the collected G. aspera propagated predominantly by sexual reproduction in Okinawan populations. The UPGMA grouping of five populations in the Okinawa, based on Nei’s unbiased genetic distance, showed two clusters that were south and north Okinawa populations. AMOVA analyses that incorporated the data of populations from the Kerama and the Ishigaki Islands detected significant FSC values among the populations within groups in analyses distinguishing north–south Okinawa, Okinawa–Kerama, and Okinawa–Kerama–Ishigaki. Significant FCT values were neither detected between north–south Okinawa nor between the Okinawa–Kerama groups. The ‘‘local-recruits’’ hypothesis, which assumes that a substantial proportion of recruits is produced locally, appears to be applicable to coral populations in the Ryukyu Archipelago. On the other hand, detection of nonsignificant FCT values indicated the presence of genetic connectivity between north–south Okinawa and Kerama–Okinawa groups. Nordemar, I. (2004) Human abuses of coral reefs – Adaptive responses and regime transitions. Stockholm University: Stockholm (Sweden), 40 pp. During the last few decades, coral reefs have become a disappearing feature of tropical marine environments, and those reefs that do remain are severely threatened. It is understood that humans have greatly altered the environment under which these ecosystems previously have thrived and evolved. Overharvesting of fish stocks, global warming and pollution are some of the most prominent threats, acting on coral reefs at several spatial and temporal scales. Presently, it is common that coral reefs have been degraded into alternative ecosystem regimes, such as macroalgae-dominated or sea urchin-barren. Although these ecosystems could

potentially return to coral dominance in a long-tem perspective, when considering current conditions, it seems likely that they will persist in their degraded states. Thus, recovery of coral reefs cannot be taken for granted on a human timescale. Multiple stressors and disturbances, which are increasingly characteristic of coral reef environments today, are believed to act synergistically and produce ecological surprises. However, current knowledge of effects of compounded disturbance and stress is limited. Based on five papers, this thesis investigates the sublethal response of multiple stressors on coral physiology, as well as the effects of compound stress and disturbance on coral reef structure and function. Adaptive responses to stress and disturbance in relation to prior experience are highlighted. The thesis further explores how inherent characteristics (traits) of corals and macroalgae may influence regime expression when faced with altered disturbance regimes, in particular overfishing, eutrophication, elevated temperature, and enhanced substrate availability. Finally, possibilities of affecting the resilience of macroalgae-dominated reefs and shifting community composition towards a coral-dominated regime are explored. Nordemar, I, Sjöö, GL, Mörk, E and McClanahan, TR. (2007) Effects of estimated herbivory on the reproductive potential of four East African algal species – a mechanism behind ecosystem shifts on coral reefs? Hydrobiologia 575, 57–68. The aim of this field study was to investigate effects of estimated fish- and sea urchin herbivory on the reproductive potential of four species of macroalgae; Halimeda macroloba (Decasine), H. renschii (Hauck), Turbinaria ornata (Turner) and Padina boergesenii (Allender et Kraft). Fish and sea urchin herbivory were calculated based on reported consumption rates for their biomass estimates. We hypothesized that reduced herbivory would increase algal size and the reproductive potential, which may promote algal recruitment and be one of the driving mechanisms behind algal shifts and persistent algae-dominated reefs. Algae were investigated in field sites where the estimated fish- and or sea urchin herbivory differed. Our results suggest that algal fecundity of T. ornata and P. boergesenii are positively correlated to their size. Fecundity of T. ornata was higher and individuals grew larger in areas where estimated fish herbivory was lower. The two species of Halimeda grew larger and had higher fecundity in areas where estimated sea urchin herbivory was lower. P. boergesenii responded ambiguously to patterns in herbivory. Due to species-specific responses to different herbivores, it is difficult to generalize about effects of overfishing on algal fecundity. Normile, D. (2000) Some coral bouncing back from El Niño. Science 288, 941-942. Coral reefs in the Indian and Pacific oceans seem to be recovering more quickly than expected from a recent devastating "bleaching" caused by high ocean temperatures. New research suggests that the nascent recoveries may be partly due to the unexpected survival of juvenile coral that somehow avoided the brunt of the environmental assault. "It may indicate that reefs are more resilient than we had thought," says Terry Done, a senior research scientist at the Australian Institute of Marine Science in Cape Ferguson who studies reefs in the Indian Ocean. However, the coral would not be able to mature and recover from the repeated bleaching forecast to accompany projected global warming, he adds.

Norström, AV, Lokrantz, J, Nyström, M and Yap, HT. (2007) Influence of dead coral substrate morphology on patterns of juvenile coral distribution. Marine Biology 150, 1145-1152. This study examines the abundances of three morphological categories of juvenile corals (massive, branching and encrusting) on two diVerent types of natural substratum, dead massive and dead branching corals. The overall results show that the morphological characteristics of dead coral substratum have a significant influence on the coral recruitment patterns with respect to the morphology of the recruits: juvenile corals of massive and branching types were more abundant on substrates of corresponding morphology. The results obtained from this study suggest that dead coral might attract coral larvae that are morphologically similar. On the other hand, it may be the result of post-settlement mortality. Whatever the mechanism shaping the patterns is, it seems that the physical morphology of the dead coral substrate has a signiWcant influence on the coral recruit assemblage. Hence, we suggest that substrate morphology can be an important qualitative factor for coral settlement and a possible determinant of community structure. Nozawa, Y, Tokeshi, M and Nojima, S. (2006) Reproduction and recruitment of scleractinian corals in a high-latitude coral community, Amakusa, southwestern Japan. Marine Biology 149, 1047–1058. Reproduction and recruitment in high-latitude coral populations in Japan have been little studied. A comprehensive study of the reproduction and early life history was conducted on nine common scleractinian coral species in Amakusa, southwestern Japan (32°N) from 2001 to 2003 including; (1) fecundity (the proportion of colonies with mature eggs), (2) timing and synchrony of spawning, (3) initial larval settlement pattern, (4) recruitment, (5) post-settlement mortality. The fecundity was high (76.7–100%) in six of seven species examined in 2002 and 2003. Annual spawning of the seven species occurred from mid July to August in 2001–2003, when seawater temperature was at the annual maximum. Spawning was highly synchronised among conspecific colonies and species in 2002 and 2003, with five species spawning five to nine nights after the full moon and another two spawning around the new moon. Temporal patterns of larval settlement of three spawning species during the first 10 days after spawning were similar to those of other spawning species from low latitudes. The number of scleractinian recruits on settlement plates, deployed from July to October (the major recruitment period at the study site), was low (2 recruits/m2) for the three consecutive years. Postsettlement mortality of 1–1.5 month old spat of five species ranged between 88 and 100% over 3–10 months in the field, similar to the values reported for both high and low latitude species (>94–99%). Among the key stages examined, the low recruitment rate may be the most important step in limiting successful reproduction and recruitment of these high-latitude scleractinian populations. The low recruitment rate may be attributable to (1) the reduced influx of larval supply from other coral populations, which are smaller and more isolated at high-latitudes and (2) the longer precompetent larval phase of broadcast-spawning corals which results in an increased chance of larvae being dispersed away from parent populations.

Nozawa, Y and Harrison, PL. (2007) Effects of elevated temperature on larval settlement and post-settlement survival in scleractinian corals, Acropora solitaryensis and Favites chinensis. Marine Biology. We examined the effects of elevated temperature under different exposure periods on larval settlement and post-settlement survival in scleractinian corals, Acropora solitaryensis and Favites chinensis. In the Wrst experiment with the subtropical coral, A. solitaryensis, the numbers of larvae settling and those dead were examined daily for 5 days at 20, 23 (ambient), 26 and 29°C conditions. Larval settlement of A. solitaryensis was initially greater at higher temperature conditions, but the peak in number of settled larvae shifted from 29 to 26°C by day 5, due to ca. 90% post-settlement mortality at 29°C condition. In order to determine the eVects under short-term exposure, larvae of F. chinensis were exposed to 27 (ambient), 31 or 34°C only for one hour in the second experiment. The number of larvae settling for 24 h after the exposure and their survivorship over subsequent week was monitored in the ambient temperature condition. Larvae of F. chinensis exhibited greater settlement at higher temperature treatments and constantly low post-settlement mortalities (< ca. 17%) in all temperature treatments, resulting in the highest number of settled larvae at 34°C treatment. These results suggested two different effects of elevated temperature on the early stages of recruitment process of scleractinian corals; (1) the positive effect on larval settlement and (2) the negative effect on post-settlement survival under prolonged exposure. Nugues, MM and Szmant, AM. (2006) Coral settlement onto Halimeda opuntia: a fatal attraction to an ephemeral substrate? Coral Reefs 25, 585-591. Degraded reefs with a high abundance of macroalgae usually also have low densities of coral recruits. Few studies, however, have examined whether these algae aVect coral larval settlement. This study demonstrates, experimentally, that larvae of the Caribbean coral Favia fragrum can settle on the green alga Halimeda opuntia even when another substrate more suitable for settlement is present. Larval settlement onto experimental substrates was quantified under three treatments: rubble only, rubble plus plastic algal mimic, and rubble plus live H. opuntia. Similar total larval settlement was observed in all treatments. No larvae settled on the algal mimic, but total settlement was similar on the rubble in the first two treatments, showing that the rubble alone offered sufficient substrate for high settlement success. About half the larvae in the live algal treatment settled on H. opuntia instead of on the rubble, showing that larvae did not reject this substrate as they did the algal mimic. This result raises the possibility that corals will settle on some macroalgae when their abundance is high. Most macroalgae, including H. opuntia, are ephemeral substrates unsuitable for post-settlement survival. Such unexpected settlement may therefore have significant consequences for coral recruitment success on algaldominated reefs.

Nyström, M, Folke, C and Moberg, F. (2000) Coral reef disturbance and resilience in a human-dominated environment. Trends in Ecology and Evolution 15, 413-417. Facing a human-dominated world, ecologists are now reconsidering the role of disturbance for coral reef ecosystem dynamics. Human activities alter the natural disturbance regimes of coral reefs by transforming pulse events into persistent disturbance or even chronic stress, by introducing new disturbance, or by suppressing or removing disturbance. Adding these alterations to natural disturbance regimes will probably result in unknown synergistic effects. Simultaneously, humans are altering the capacity of reefs to cope with disturbance (e.g. by habitat fragmentation and reduction of functional diversity), which further exacerbates the effects of altered disturbance regimes. A disturbance that previously triggered the renewal and development of reefs might, under such circumstances, become an obstacle to development. The implications of these changes for reef-associated human activities, such as fishing and tourism, can be substantial. Nyström, M and Folke, C. (2001) Spatial resilience of coral reefs. Ecosystems 4, 406-417. There have been several earlier studies that addressed the influence of natural disturbance regimes on coral reefs. Humans alter natural disturbance regimes, introduce new stressors, and modify background conditions of reefs. We focus on how coral reef ecosystems relate to disturbance in an increasingly human-dominated environment. The concept of ecosystem resilience—that is, the capacity of complex systems with multiple stable states to absorb disturbance, reorganize, and adapt to change—is central in this context. Instead of focusing on the recovery of certain species and populations within disturbed sites of individual reefs, we address spatial resilience—that is, the dynamic capacity of a reef matrix to reorganize and maintain ecosystem function following disturbance. The interplay between disturbance and ecosystem resilience is highlighted. We begin the identification of spatial sources of resilience in dynamic seascapes and exemplify and discuss the relation between “ecological memory” (biological legacies, mobile link species, and support areas) and functional diversity for seascape resilience. Managing for resilience in dynamic seascapes not only enhances the likelihood of conserving coral reefs, it also provides insurance to society by sustaining essential ecosystem services. Nyström, M. (2006) Redundancy and response diversity of functional groups: Implications for the resilience of coral reefs. Ambio 35, 30-35. To improve coral reef management, a deeper understanding of biodiversity across scales in the context of functional groups is required. The focus of this paper is on the role of diversity within functional groups in securing important ecosystem processes that contribute to the resilience of coral-dominated reef states. Two important components of species biodiversity that confer ecosystem resilience are analyzed: redundancy and the diversity of responses within functional groups to change. Three critical functional groups are used to illustrate the interaction between these two components and their role in coral reef resilience: zooxanthellae (symbiotic micro algae in reef-building corals), reef-building corals, and herbivores. The paper further examines the consequences of undermining functional redundancy and response

diversity and addresses strategies to secure ecological processes that are critical for coral reef resilience. Obura, DO. (2001) Can differential bleaching and mortality among coral species offer useful indicators for assessment and management of reefs under stress? Bulletin of Marine Science 69, 421-442. Coral reefs are increasingly subject to a variety of threats from local to global scales. The development of monitoring and assessment tools to diagnose coral reef community responses to threats would assist greatly in management of their impacts. This paper examines interspecific patterns of bleaching, mortality and recovery of corals on reefs in Kenya, Tanzania, Mozambique and Madagascar for one year after the 1997-98 El Niño. On average, 50-80% bleaching and mortality of corals occurred throughout the region. Coral tissue condition (normal, pale, bleached and dead) was recorded for up to one year after the onset of bleaching. Thirteen coral species were used for analysis. Cluster Analysis and Principal Components Analysis with factor rotation were applied to the dataset, giving 2 major groups of species, characterized by a) severe bleaching followed by 100% mortality (e.g. Acropora spp., Pocillopora spp., Galaxea astreata), b) graduated bleaching with pale tissue and low to moderate mortality (e.g. Porites lutea, Echinopora gemmacea, Hydnophora exesa), and one outlier with long term persistence of bleaching (Montipora tuberculosa). These species groups based on the bleaching response are consistent with life history strategies proposed by other workers. It is proposed that observations of coral condition and bleaching in the field, interpreted in the light of life history theory, can provide a coral bioassay technique for reef monitoring and management. Obura, D. (2001) Differential bleaching and mortality of eastern African corals. In Richmond, MD and Francis, J (eds.) Marine science development in Tanzania and Eastern Africa: Proceedings of the 20th Anniversary Conference on Advances in Marine Sciences in Tanzania. IMS/WIOMSA, 301-317. Coral reefs are increasingly subject to a variety of threats both at the local and global scale. The development of monitoring and assessment tools to diagnose coral reef community responses to threats would assist greatly in the management of their impacts. This paper examines interspecific patterns of bleaching, mortality and recovery of corals on reefs in Kenya, Tanzania, Mozambique and Madagascar for one year after the 1997–98 El Niño weather phenomenon. On average, 50–80% bleaching of corals was observed, with a similar overall proportion suffering mortality. Coral tissue condition (normal, pale, bleached and dead) was recorded for up to one year after the onset of bleaching. Seventeen coral species had sufficient numbers of colonies in each time period for analysis. Cluster analysis and principal components analysis with factor rotation were applied to the dataset, giving 3 major groups of species, characterised by (a) severe bleaching followed by 100% mortality (seen in Acropora spp., Pocillopora spp., Galaxea astreata), (b) graduated bleaching with pale tissue with low to moderate mortality (Porites lutea, Echinopora gemmacea, Hydnophora exesa), and (c) moderate, persistent bleaching with moderate recovery (Pavona varians, Montipora tuberculosa). These species groups based on the bleaching response are consistent with coral life history strategies proposed by other workers.

Obura, D and Mangubhai, S. (2001) Assessing environmental and ecological factors and their contributions to coral bleaching resistance and resilience of reefs in the Western Indian Ocean. Proceedings of the International Coral Reef Initiative Regional Workshop for the Indian Ocean: Maputo (Mozambique), 15 pp. Large-scale coral bleaching events have increased in intensity, frequency, and geographic distribution in the last two decades, caused by increasingly severe El Niño Southern Oscillation (ENSO) events. However, despite the widespread bleaching associated mortality of coral reef organisms, it is rare for living corals to be completely eliminated from a section of reef. Even in the most severe cases, some coral communities appear to be more resistant (coral colonies do not bleach or bleach but do not die) or more resilient (the colonies bleach and partially or entirely die but recover rapidly for a variety of reasons) to stresses that cause bleaching. A questionnaire designed to investigate the association between different resistance and resilience factors was developed in an initiative spearheaded by The Nature Conservancy and the World Wildlife Fund. This study analyses preliminary results based on circulation of the questionnaire in the western Indian Ocean, where coral reefs experienced wide-scale coral bleaching during the 1998 El Niño event. The findings of this study indicate that depth and steep reef slopes provided protection from coral bleaching, but a variety of other factors expected to do the same, such as high levels of temperature fluctuations, high levels of turbidity, and a robust reef community did not. Potential interactions and complications in the dataset are discussed. The results will offer some guidance on the potential to use environmental variables to develop guidelines on how existing management can be designed to mitigate the impacts of climate change, such as by focussing on reefs that may act as refugia during bleaching events, and/or which support the recovery of adjacent impacted or degraded reefs. Obura, D. (2005) Resilience and climate change: Lessons from coral reefs and bleaching in the Western Indian Ocean. Estuarine, Coastal and Shelf Science 63, 353-372. The impact of climate change through thermal stress-related coral bleaching on coral reefs of the Western Indian Ocean has been well documented and is caused by rising sea water temperatures associated with background warming trends and extreme climate events. Recent studies have identified a number of factors that may reduce the impact of coral bleaching and mortality at a reef or subreef level. However, there is little scientific consensus as yet, and it is unclear how well current science supports the immediate needs of management responses to climate change. This paper provides evidence from the Western Indian Ocean in support of recent hypotheses on coral and reef vulnerability to thermal stress that have been loosely termed ‘resistance and resilience to bleaching’. The paper argues for a more explicit definition of terms, and identifies three concepts affecting coral-zooxanthellae holobiont and reef vulnerability to thermal stress previously termed ‘resistance to bleaching’: ‘thermal protection’, where some reefs are protected from the thermal conditions that induce bleaching and/or where local physical conditions reduce bleaching and mortality levels; ‘thermal resistance’, where individual corals bleach to differing degrees to the same thermal stress; and ‘thermal tolerance’, where individual corals suffer differing levels of mortality when exposed to the same thermal

stress. ‘Resilience to bleaching’ is a special case of ecological resilience, where recovery following large-scale bleaching mortality varies according to ecological and other processes. These concepts apply across multiple levels of biological organization and temporal and spatial scales. Thermal resistance and tolerance are genetic properties and may interact with environmental protection properties resulting in phenotypic variation in bleaching and mortality of corals. The presence or absence of human threats and varying levels of reef management may alter the influence of the above factors, particularly through their impacts on resilience, offering the opportunity for management interventions to mitigate the impacts of thermal stress and recovery on coral reefs. These concepts are compiled within an overarching framework of spatial resilience theory. This provides a framework for developing linked scientific and management questions relating to the larger scale impacts of climate change on coral reefs, their management needs and prospects for their future. Okamoto, M, Nojima, S, Furushima, Y and Phoel, WC. (2005) A basic experiment of coral culture using sexual reproduction in the open sea. Fisheries Science 71, 263-270. Coral larvae, produced from a mass spawning event, were successfully settled on special stone settlement sticks and raised in situ for eventual transport to other reefs. The test area, Sekisei lagoon, Okinawa Prefecture, Japan, is located close to the warm Kuroshio current and is surmised to be the source from which major Japanese corals are derived. A total of 131 settlement sticks, with small holes in their sides to increase protection from grazing (4 mm in diameter and approximately 5 mm deep), were deployed in the lagoon the day before the coral’s mass spawning. After 3 months, 61 sticks were recovered containing 71 corals, mostly in the holes. After 1 year, three corals were confirmed to be growing well and extending outside the holes of the three sticks out of 70 sticks left in the water at the lagoon site. They survived two potentially lethal conditions, that is, high water temperatures with associated extensive coral bleaching and continuous grazing pressure from predators. This procedure is applicable for large-scale coral transplantation, not only in Japan but also in other tropical countries. Okubo, N, Taniguchi, H and Motokawa, T. (2005) Successful methods for transplanting fragments of Acropora Formosa and Acropora hyacinthus. Coral Reefs 24, 333-342. In order to establish a successful method for the transplantation of branching and tabular coral fragments, we tested the effects of orientations of attachment, seasons of transplantation, and size of fragments on survival, growth, and spawning using Acropora formosa and A. hyacinthus. Vertically attached, large-sized fragments of A. formosa showed 98–100% survival rate after 18 months. The fragments transplanted in August exhibited better survival than those transplanted in November. The larger fragments had the higher percentage of spawning. The fragments that spawned had lower growth rate, while those resorbed the oocytes carried at the time of transplantation showed higher growth rate, suggesting the trade-off between growth and reproduction. Half of the fragments spawned 1 month earlier than the donor colonies. Only the vertically attached fragments of A. hyacinthus fused to the

substratum, and those transplanted in February showed 100% survival rate after 14 months, indicating that this species is well suited for transplantation. Oliver, J, Marshall, P, Setiasih, N and Hansen, L. (2004) A global protocol for assessment and monitoring of coral bleaching. WorldFish Centre, Penang (Malaysia), 35 pp. Coral bleaching occurs when corals lose the single celled algae (zooxanthellae) which live within their tissues. These golden brown coloured algae (Figure 1) occur in varying densities in reef corals (and other reef invertebrates), and give them a light tan to deep chocolate brown colour. Where additional pigments exist within the animal cells, this brown colour can be overlain by different additional hues such as blue, green, purple or yellow (Figure 2). When corals lose their zooxanthellae, the white skeleton can be seen through the transparent animal tissue, making the corals looked bleached white. In cases where the corals possess additional animal pigments, bleached corals take on vivid fluorescent hues, with no trace of the normal brown background colour (Figure 3). Corals which have bleached are not dead. On close inspection it is possible to see the transparent polyps and tentacles of bleached colonies. However corals cannot remain bleached indefinitely. If the stress is not too severe or prolonged, stressed colonies can slowly regain their zooxanthellae and survive. But in severe bleaching events many corals subsequently die, casing major changes to the structure and function of the reef ecosystem, and possible cascading impacts on other organisms. Although bleaching in individual corals was first observed early in this century, massive events affecting reefs spanning tens to hundreds of kilometres have only been documented in the last two decades . In the laboratory, or on a small scale in the field, bleaching has been attributed to a number of factors which cause stress, including extremes (both high and low) of temperature and light, low salinity, low oxygen, high concentration of toxic chemicals which affect respiration and photosynthesis. In the case of “mass coral bleaching” involving a range of coral species over large areas, elevated temperatures are the primary stress factor (Coles & Brown, 2003; Hoegh-Guldberg 1999). Elevated sea temperatures may be driven by regional phenomena (such as El Niño events), but local conditions, such as periods of calm weather and clear skies, also play a major role in determining sea temperatures in reefal regions. Decreased salinity from excessive rainfall and flood plumes can increase the amount of bleaching, and may be a primary cause in some situations (mostly over limited spatial scales). During the worst global bleaching event in 1998, more than 680 records of coral bleaching were reported from over 55 different reef regions. Many of the affected reefs suffered massive mortality from which they are still recovering (Wilkinson 2002). This event was also the most intensively studied, and satellite data showing thermal anomalies or “hot spots” clearly showed that large patterns of abnormally and consistently high temperature were very well correlated with bleaching patterns, and in some cases appeared to be able to predict the events (Toscano et al. 2000, Wilkinson et al 1999). Coral bleaching is now considered to be one of the most significant and widespread threats to coral reefs Hughes et al., 2003). When predicted temperature increases due to global warming over the next 100 years have been compared to the known temperature bleaching limits of corals, the depressing conclusion is that by 2020 coral reefs in many parts of the world may suffer bleaching and mortality every year Hoegh-Guldberg, 1999). Unless corals are able to adapt to raising temperatures,

reefs may suffer progressive deterioration and species loss, resulting in major ecological impacts and consequent social and economic impacts on the human communities in many countries that depend on reefs for their livelihoods. Global warming and greenhouse gas emission thus represent a relatively new and severe threat to the sustainability and productivity of coral reefs and the services that that provide to humans (Hughes et al., 2003; Wellington et al., 2001; Cesar et al., 2003). There is an urgent need to obtain better information on bleaching events around the world. This information is crucial to scientific understanding of the fate of coral reefs and to the feasibility and practicality of developing management strategies to increase the resistance and resilience of reefs to bleaching and associated mortality events. This protocol aims to provide a simple yet consistent set of procedures to: 1. Document the spatial distribution, timing and severity of large scale bleaching events and how they compare with maps of global temperature anomalies and other meteorological and oceanographic factors. This will help in the developing and testing of predictive models of bleaching which can focus attention on susceptible reefs which might benefit from better management of additional human impacts, and identify reefs which need to be protected since they are most likely to survive further warming and bleaching events. 2. Determine the factors that, in addition to increased sea temperature, play a major role in increasing (and decreasing) susceptibility to bleaching and subsequent mortality. 3. Develop a better understanding of which existing and new management practices can help reefs deal with the additional stresses of a warming ocean. 4. Raise awareness of the extent and urgency of coral bleaching as a possible threat to the future health and existence of coral reefs. Olsson, P, Folke, C and Berker, F. (2003) Adaptive co-management for building resilience in social-ecological systems. Stockholm University: Stockholm (Sweden), 48 pp. Ecosystems are complex adaptive systems that require flexible governance with the ability to respond to environmental feedback. We present, through examples from Sweden and Canada, the development of adaptive co-management systems, showing how local groups self-organize, learn and actively adapt to and shape change with social networks that connect institutions and organizations across levels and scales and that facilitate information flows. The development took place through a sequence of responses to environmental events that widened the scope of local management from a particular issue or resource to a broad set of issues related to ecosystems processes across scales and from individual actors, to group of actors to multiple-actor processes. The results suggest that the institutional and organizational landscape should be approached as carefully as the ecological in order to clarify features that contribute to the resilience of social-ecological systems. These include vision, leadership and trust; enabling legislation that creates social space for ecosystem management; funds for responding to environmental change and for remedial action; capacity for monitoring and responding to environmental feedback; information flow through social networks; the combination of various sources of information and knowledge, sense-making and arenas of collaborative learning for ecosystem management. We conclude that the self-organizing process of adaptive co-management development, facilitated by rules and incentives of higher levels, has

the potential to expand desirable stability domains of a region and make social-ecological systems more robust to change. Omori, M, Fukami, H, Kobinata, H and Hatta, M. (2001) Significant drop of fertilization of Acropora corals in 1999: An after-effect of heavy coral bleaching? Limnology and Oceanography 46, 704-706. In June 1999, after devastating coral bleaching in 1998, laboratory fertilization of Acropora nasuta, one of the most abundant reef-building corals in Okinawa, Japan, decreased significantly from usual rates (>94%) to an average of 42% at a sperm concentration of 105 ml-1, Similar decreases were observed in four other mass-spawning acroporid corals. We also found a decrease in sperm motility in the laboratory. A series of experiments to determine the effects of sperm concentration on fertilization rates revealed that sperm of 107 ml-1 was needed to obtain a rate >80%. Sperm concentration in surface seawater during mass spawning was highest within 1.0 it of spawning but decreased sharply thereafter. These results suggest that gamete dilution plays an important role in limiting the fertilization of coral eggs in the sea. As successful fertilization appears to have been much lower in 1999, we suspect that production of new coral recruits was also reduced greatly. Current and future sea-temperature increases thus pose a severe potential threat to coral reefs by increasing the frequency of coral bleaching and consequently leading to further declines of coral recruitment and hence, reef corals. Ostrander, GK, Armstrong, KM, Knobbe, ET, Gerace, D and Scully, EP. (2000) Rapid transition in the structure of a coral reef community: The effects of coral bleaching and physical disturbance. Proceedings of the National Academy of Sciences of the United States of America 97, 5297-5302. Coral reef communities are in a state of change throughout their geographical range. Factors contributing to this change include bleaching (the loss of algal symbionts), storm damage, disease, and increasing abundance of macroalgae. An additional factor for Caribbean reefs is the aftereffects of the epizootic that reduced the abundance of the herbivorous sea urchin, Diadema antillarum. Although coral reef communities have undergone phase shifts, there are few studies that document the details of such transitions. We report the results of a 40-month study that documents changes in a Caribbean reef community affected by bleaching, hurricane damage, and an increasing abundance of macroalgae. The study site was in a relatively pristine area of the reef surrounding the island of San Salvador in the Bahamas. Ten transects were sampled every 3-9 months from November 1994 to February 1998. During this period, the corals experienced a massive bleaching event resulting in a significant decline in coral abundance. Algae, especially macroalgae, increased in abundance until they effectively dominated the substrate. The direct impact of Hurricane Lili in October 1996 did not alter the developing community structure and may have facilitated increasing algal abundance. The results of this study document the rapid transition of this reef community from one in which corals and algae were codominant to a community dominated by macroalgae. The relatively brief time period required for this transition illustrates the dynamic nature of reef communities.

Otis, DB, Carder, KL, English, DC and Ivey, JE. (2004) CDOM transport from the Bahamas Banks. Coral Reefs 23, 152-160. The transport of colored dissolved organic matter (CDOM) between shallow banks and deep basins in the Bahamas was the focus of this study. Hydrographic and CDOM absorption measurements made on the Bahamas Banks and in Exuma Sound during the spring of 1999 and 2000 showed that values of salinity and CDOM absorption at 440 nm were higher on the banks (37.18 psu, 0.06 m–1), compared to Exuma Sound (37.04 psu, 0.03 m–1). Spatial patterns of CDOM absorption in Exuma Sound revealed that plumes of CDOM-rich water flow into Exuma Sound from the surrounding banks. These patterns were determined using Sea-viewing Wide Field-of-view Sensor (SeaWiFS) data processed using a Moderate Resolution Imaging Spectroradiometer (MODIS) algorithm to derive CDOM absorption estimates. These data, along with time-series data collected in a channel between the banks and sound, suggest that bank water rich in CDOM and salinity leaves the banks during ebb tide, whereas sound water, with lower levels of CDOM and salinity, extends onto the banks during flood tide. Because CDOM absorbs ultraviolet radiation, a causal factor of reef organism bleaching, we discuss the meaning of our findings in terms of susceptibility to coral bleaching in the Exuma region. Paddack, MJ, Cowen, RK and Sponaugle, S. (2006) Grazing pressure of herbivorous coral reef fishes on low coral-cover reefs. Coral Reefs 25, 461–472. The impact of grazing by herbivorous fishes (Acanthuridae, Scaridae, and Pomacentridae) on low coral-cover reefs was assessed by measuring rates of benthic algal production and consumption on inshore and offshore reefs in the upper Florida Keys. Algal production rates, determined in situ with caged and uncaged experimental plates, were low (mean 1.05 g C m-2 day-1) and similar among reef types. Algal consumption rates were estimated using two different models, a detailed model incorporating fish bite rates and algal yield-per-bite for one species extrapolated to a guild-wide value, and a general regression relating fish biomass to algal consumption. Algal consumption differed among reef types: a majority of algal production was consumed on offshore reefs (55–100%), whereas consumption on inshore patch reefs was 31–51%. Spatial variation in algal consumption was driven by differences in herbivorous fish species composition, density, and size-structure among reef types. Algal consumption rates also varied temporally due to seasonal declines in bite rates and intermittent presence of large-bodied, vagile, schooling species. Spatial coherence of benthic community structure and temporal stability of algal turf over 3 years suggests that grazing intensity is currently sufficient to limit further spread of macroalgal cover on these low coral-cover reefs, but not to exclude it from the system. Palumbi, SR. (2002) Marine reserves: A tool for ecosystem management and conservation. Pew Oceans Commission: Arlington (USA), 44 pp. Abstract not available.

Palumbi, SR. (2003) Population genetics, demographic connectivity, and the design of marine reserves. Ecological Applications 13, S146-S158. Genetic analyses of marine population structure often find only slight geographic differentiation in species with high dispersal potential. Interpreting the significance of this slight genetic signal has been difficult because even mild genetic structure implies very limited demographic exchange between populations, but slight differentiation could also be due to sampling error. Examination of genetic isolation by distance, in which close populations are more similar than distant ones, has the potential to increase confidence in the significance of slight genetic differentiation. Simulations of one-dimensional stepping stone populations with particular larval dispersal regimes shows that isolation by distance is most obvious when comparing populations separated by 2–5 times the mean larval dispersal distance. Available data on fish and invertebrates can be calibrated with this simulation approach and suggest mean dispersal distances of 25–150 km. Design of marine reserve systems requires an understanding of larval transport in and out of reserves, whether reserves will be self-seeding, whether they will accumulate recruits from surrounding exploited areas, and whether reserve networks can exchange recruits. Direct measurements of mean larval dispersal are needed to understand connectivity in a reserve system, but such measurements are extremely difficult. Genetic patterns of isolation by distance have the potential to add to direct measurement of larval dispersal distance and can help set the appropriate geographic scales on which marine reserve systems will function well. Pandolfi, JM, Bradbury, RH, Sala, E, Hughes, TP, Bjorndal, KA, Cooke, RG, McArdle, D, McClenachan, L, Newman, MJH, Paredes, G, Warner, RR and Jackson, JBC. (2003) Global trajectories of the long-term decline of coral reef ecosystems. Science 301, 955-958. Degradation of coral reef ecosystems began centuries ago, but there is no global summary of the magnitude of change. We compiled records, extending back thousands of years, of the status and trends of seven major guilds of carnivores, herbivores, and architectural species from 14 regions. Large animals declined before small animals and architectural species, and Atlantic reefs declined before reefs in the Red Sea and Australia, but the trajectories of decline were markedly similar worldwide. All reefs were substantially degraded long before outbreaks of coral disease and bleaching. Regardless of these new threats, reefs will not survive without immediate protection from human exploitation over large spatial scales. Pante, E, King, A and Dustan, P. (2007) Short-term decline of a Bahamian patch reef coral community: Rainbow Gardens Reef 1991–2004. Hydrobiologia. Temporal changes in the stony coral assemblages of RainbowGardens Reef (Iguana Cay, Exumas, Bahamas), were revealed by comparing quantitative community descriptors recorded in 1991 and 2004. Mean percent live cover significantly dropped from 13% to 3% between 1991 and 2004 (-77%, P < 0.01). During that period, coral abundance (number of colonies) decreased from 295 to 240 (-18.6%, P < 0.01). The community was less rich, equally diverse, more even, and the spatial structure of the reef had become more homogeneous. Most large Montastrea annularis, Agaricia agaricites, Porites porites, and Porites astreoides colonies were absent in 2004.

Colonies were also more prone to stress related to algal smoothering, excess sediment, and endolithic boring. No disease was recorded on stony corals in 2004. Examination of M. annularis skeletons revealed intense bioerosion and a bright yellow skeletal band that corresponded to 1998–1999. Reefs worldwide suffered an unprecedented period of coral bleaching during the 1998 El Niño event, which is likely to have played a role in the decline of Rainbow Gardens. Evidence suggests that the synergistic actions of bleaching, bioerosion, and storm action reduced Rainbow’s formerly discrete coral patches to rubble. These results attest to the state of crisis of Caribbean coral reefs: even shallow, and presumably eurythermic communities in remote localities can be sensitive to change. Other patch reefs in the vicinity of Rainbow Gardens do not seem as degraded suggesting that local, small-scale differences may be important components of reef resilience. Parmesan, C and Yohe, G. (2003) A globally coherent fingerprint of climate change impacts across natural systems. Nature 421, 37-42. Causal attribution of recent biological trends to climate change is complicated because non-climatic influences dominate local, short-term biological changes. Any underlying signal from climate change is likely to be revealed by analyses that seek systematic trends across diverse species and geographic regions; however, debates within the Intergovernmental Panel on Climate Change (IPCC) reveal several definitions of a ‘systematic trend’. Here, we explore these differences, apply diverse analyses to more than 1,700 species, and show that recent biological trends match climate change predictions. Global meta-analyses documented significant range shifts averaging 6.1 km per decade towards the poles (or metres per decade upward), and significant mean advancement of spring events by 2.3 days per decade. We define a diagnostic fingerprint of temporal and spatial ‘sign-switching’ responses uniquely predicted by twentieth century climate trends. Among appropriate long-term/large-scale/multi-species data sets, this diagnostic fingerprint was found for 279 species. This suite of analyses generates ‘very high confidence’ (as laid down by the IPCC) that climate change is already affecting living systems. Parmesan, C. (2006) Ecological and Evolutionary Responses to Recent Climate Change. Annual Review of Ecology and Evolution and Systematics. 37, 637–69. Ecological changes in the phenology and distribution of plants and animals are occurring in all well-studied marine, freshwater, and terrestrial groups. These observed changes are heavily biased in the directions predicted from global warming and have been linked to local or regional climate change through correlations between climate and biological variation, field and laboratory experiments, and physiological research. Range-restricted species, particularly polar and mountaintop species, show severe range contractions and have been the first groups in which entire species have gone extinct due to recent climate change. Tropical coral reefs and amphibians have been most negatively affected. Predator-prey and plant-insect interactions have been disrupted when interacting species have responded differently to warming. Evolutionary adaptations to warmer conditions have occurred in the interiors of species’ ranges, and resource use and dispersal have evolved rapidly at expanding range margins. Observed genetic shifts modulate local effects of climate change, but there is little evidence that they will mitigate negative effects at the species level.

Peterson, G, Allen, CR and Holling, CS. (1998) Ecological resilience, biodiversity, and scale. Ecosystems 1, 6-18. We describe existing models of the relationship between species diversity and ecological function, and propose a conceptual model that relates species richness, ecological resilience, and scale. We suggest that species interact with scale-dependent sets of ecological structures and processes that determine functional opportunities. We propose that ecological resilience is generated by diverse, but overlapping, function within a scale and by apparently redundant species that operate at different scales, thereby reinforcing function across scales. The distribution of functional diversity within and across scales enables regeneration and renewal to occur following ecological disruption over a wide range of scales. Pet-Soede, L. (2000) Effects of coral bleaching on the socio-economics of the fishery in Bolinao, Pangasinan, Philippines. Coastal Resources Center University of Rhode Island: Narragansett (USA), 23 pp. Socio-economic effects of bleaching, if any, were estimated from literature, interviews and catch logbooks that were also used in a cost-benefit analysis of the fishery at the household level. Trends in total catch and in Catch per unit Effort (CpuE) were studied for effecst of the 1998 bleaching event from catch and effort data that were collected since 1996 by the Marine Fisheries Resources Management Project (MFRMP). Perceptions of fishers on causes to changes in their fishery and on the importance of bleaching were inventoried. The status of the fishery was compared to recent changes in habitat and fish community structure. An estimated total of 3154 fishers are engaged in the fishery at and around the Bolinao reef area (200 km2). A large variety of fishing gears is used and patterns in resource utilisation vary with different requirements for operation of the gear and differences in target species. Of the reef fishery gears, large-scale spear gun fishers using compressors catch most, followed by triplet nets. The parisris and deep-sea hook and line are most efficient catching some 6-8 kg of fish per hr. Lowest catch rates of 0.25-1 kg per hr are found for gillnets that are operated during the day, small-scale spear guns and hook and line. Logbooks confirm these average catches and show the uncertainty of the fisheries indicated by large daily variances around the average catches. The average monthly income in 1985 for small-scale Bolinao fishers was estimated at some 470 P per month with household expenditures almost doubling the income and it was concluded that fishing does not support minimum financial needs for survival. At present however, it appears that each fisher is able to feed his family and save some money for other expenses. Prices vary at the different points in the product chain and increase sometimes as much as 300% from one level to the next. The costs of operation varies between fisheries with highest total costs for larger boats and when using expensive technology. When we use our estimates for revenue and costs average gross income for the different fisheries varied between 125 – 300 US$. For an estimate of the added value of a fishery, the opportunity costs of labour are deducted. The result shows that except for the shell collector, who would not consider another occupation as they are mostly women and children, every operation makes significantly more money fishing than if they were working as an unskilled labourer. Net incomes after deducting opportunity costs of labour vary between 680- 8840 P per month. Especially the larger scale operations, even after dividing the income over crewmembers, have significant monthly incomes. Fishers using crab

pots and lagrite have lowest net incomes but these have often other (fishery) sources of income. There appears little difference between catch rates in 1997 (before bleaching) and in 1999 (after bleaching) for most fishing gears. Only catch rates for tabar or lambat, a shallow water gill net operated while water beating, are clearly higher than before 1999. These fishers aim mostly at herbivores such as rabbitfish and parrotfish. Even when accounting for the level of fishing effort, again only tabar nets show a significant increase in CpUE at an even slightly higher level of fishing effort. Thus it appears that only this fishery benefited from changes that occurred in 1998. This could be related to the bleaching event, nevertheless from the underwater visual census data we cannot find sufficient arguments for particular shifts in the fish community that would only affect this particular fishery. When asked, most of the gill net fishers actually said that their catches had reduced over the past years, no one mentioned increasing catches. From the interviews we found that 57% of the fishers had heard about bleaching and 53% had witnessed it personally. Of the people who had heard about or had witnessed the bleaching, 79% said that it was caused by the use of sodium cyanide in the illegal fishery for aquarium and live food fish. Only 21% said that it was caused by El Niño and most of these had probably heard the correct explanation of the phenomenon during community consultation meetings. Some thought that it caused catches to drop. When asked about their catches, 90% of the fishers said that the catch biomass had changed since 5 years ago. 8% said there is more fish now as a result of better management (stricter enforcement of the ban on destructive fishing) and 89% said that the catches had gone down mostly because of over fishing (64%) and destructive fishing (14%). Of all fishers 33% said that size in the catch has also changed, and is smaller (82%) or larger (18%) than 5 years ago. Of all fishers interviewed 43% said that the species composition has changed, and that some species cannot be caught anymore (such as cardinalfish). Most fishers (37%) suggested that strict enforcement of the recently established fishery ordinance was the best solution. Underwater visual census data indicates that the habitat structure at Bolinao reefs has changed drastically and that this is significantly related to the 1998 bleaching event. Changes in the fish community structure from 1996 to 1999 cannot be so easily explained. Fish abundance and biomass increased after 1996 but this trend didn’t change after the bleaching. Trends in catches show no clear change in the year after the bleaching except for increased catches with the small-scale gill net called tabar. Only if the 100% increase in catches of tabar fishers is indeed related to the bleaching it could be argued that the bleaching affected their income positively. None of the fishers that use this gear mentioned this increase however. It can also not be fully explained from changes in the reef fish community structure, because then other fishing gears should catch more fish as well. Therefore it is concluded from the available data that there is no evident effect of the 1998-bleaching event on the fishery in Bolinao as of yet. The fact that Bolinao fishers appear to be depending on herbivorous fish that fetch a relatively high price at the local market presents a situation that is interesting from a scientific viewpoint. In theory the Bolinao fishers could be better off after a bleaching event, when the herbivorous fish manage to benefit from the extended algal growth following coral mortality. Nevertheless, regardless effects of bleaching, this is a clear example of a disrupted system with little scope for a healthy reef fishery.

Pomeroy, RS, Parks, JE and Watson, LM. (2004) How is your MPA doing? A guidebook of natural and social indicators for evaluating Marine Protected Areas. IUCN: Gland (Switzerland), 216 pp. Effective management of marine protected areas (MPAs) requires continuous feedback of information to achieve objectives. In 2000, a collaborative initiative was launched to improve the management of MPAs. The initiative focused on working with managers, planners, and other decision-makers to develop a set of indicators for assessing the effectiveness of MPA use. This initiative was aimed at both enhancing the potential and capability for adaptive management of MPAs, as well as improving our understanding of how effective MPAs that are now being used around the world. This paper presents an overview of the MPA management effectiveness methodology and indicators, summary results of the testing phase of the guidebook at 18 MPAs around the world, and considerations for its application and use as an adaptive management tool for MPAs. Pratchett, MS, Wilson, SK, Berumen, ML and McCormick, MI. (2004) Sublethal effects of coral bleaching on an obligate coral feeding butterflyfish. Coral Reefs 23, 352-356. Coral bleaching is a significant and increasingly prevalent source of coral mortality, representing one of the most severe and widespread disturbances affecting coral reef ecosystems (Hoegh-Guldberg 1999; Pockley 2000). In the last few years (mostly since 1998), major episodes of coral bleaching have occurred on many coral reefs throughout the world, killing 20–80% of zooxanthellate corals (including both scleractinians and alcyonaceans) across expansive reef areas (e.g., Great Barrier Reef, Baird and Marshall 1998; Japan, Shibuno et al. 1999; eastern Pacific, Glynn et al. 2001; Caribbean, Ostrander et al. 2000). In addition to killing zooxanthellate corals, severe large-scale bleaching events may cause significant declines in the abundance of coral reef fishes, particularly among reef fish species that depend on live coral for food or shelter (Shibuno et al. 1999; Kokita and Nakazono 2001; Adjeroud et al. 2002). Among those fishes with the greatest reliance on live corals are butterflyfish from the genus Chaetodon (family Chaetodontidae), most of which feed primarily (if not exclusively) on living tissue from scleractinian and alcyonacean corals (e.g., Birkeland and Neudecker 1981; Harmelin-Vivien and Bouchan-Navaro 1983; Anderson et al. 1981). Further, spatial and temporal variation in the abundance of Chaetodon butterflyfish is often associated with variation in the abundance of live corals (Findley and Findley 1985). Most notably, significant declines in the abundance of Chaetodon butterflyfish have been observed following coral depletion by crown-of-thorns starfish (Bouchan-Navaro et al. 1985; Williams 1986; Sano et al. 1987). Therefore, similar declines would be expected where bleaching causes extensive coral mortality. Even if there is no immediate decline in the abundance of butterflyfish, the reduced availability of prey corals is likely to affect the physiological condition and subsequent fitness of coral-feeding butterflyfish (Jones and McCormick 2002). In this study, we explored the effects of coral bleaching on the redfin butterflyfish, Chaetodon lunulatus Quoy and Gaimard 1824. This species is distributed throughout the Indo-Pacific Archipelago and eastern Pacific Ocean (Allen et al. 1998), and is among the most abundant butterflyfish species on reefs within this region (e.g.,

Anderson et al. 1981; Bouchan-Navaro et al. 1985). Previous studies have shown that Chaetodon lunulatus feeds exclusively on living tissues from scleractinian corals (Harmelin-Vivien and Bouchan-Navaro 1983), and is likely therefore, to be very susceptible to changes in the availability of scleractinian corals (Kokita and Nakazono 2001). The purpose of this study was to investigate changes in the (1) abundance, (2) dietary habits (specifically, the proportional consumption of different prey corals), and (3) physiological condition of C. lunulatus, during a major episode of coral bleaching. The specific effects of coral bleaching on C. lunulatus were investigated over a 2-year period (May 2000 to March 2002), on the Great Barrier Reef (GBR), Australia. During this period, the GBR was subject to the most severe episode of coral bleaching ever recorded (Dennis 2002). Pratchett, MS, Wilson, SK and Baird, AH. (2006) Declines in the abundance of Chaetodon butterflyfishes following extensive coral depletion. Journal of Fish Biology 69, 1269–1280. This study documented temporal variation in the abundance of butterflyfishes (Chaetodontidae) at Trunk Reef, on the central Great Barrier Reef, Australia, from May 2000 to March 2005. During this period, live coral cover declined by >90%, mostly due to severe coral bleaching. There were no short-term changes (within 4 months) in the abundance of butterflyfishes following initial declines in live coral cover. Surveys conducted in 2005, however, revealed significant declines in the abundance of Chaetodon baronessa, Chaetodon lunulatus, Chaetodon trifascialis, Chaetodon plebeius and Chaetodon rainfordi, all of which are obligate hard-coral feeders. In contrast, there was no significant change in the abundance of Chaetodon auriga, Chaetodon aureofasciatus, Chaetodon citrinellus, Chaetodon melannotus or Chaetodon vagabundus, which are much less reliant on scleractinian coral for food. Clearly, extensive coral depletion, such as that caused by severe coral bleaching, can have a major effect on the abundance of butterflyfishes. Specific responses of butterflyfishes varied according to their reliance on hard corals for food and their ability to utilize alternate prey types. Precht, WF and Aronson, RB. (2004) Climate flickers and range shifts of reef corals. Frontiers in Ecology and Environment 2, 307-314. Staghorn coral (Acropora cervicornis) and elkhorn coral (Acropora palmata), are important reef builders in the Caribbean. In the early to middle Holocene (10 000–6000 years ago), when sea temperatures were warmer than today, Acropora-dominated reefs were common along the east coast of Florida as far north as Palm Beach County. The fossil record shows that the northern limits of these two cold sensitive species subsequently contracted to Biscayne Bay, south of Miami, apparently as a result of climatic cooling. This response of the Acropora species to climate provides a context for interpreting recent shifts in their geographic distribution. Despite recent disease-induced mass mortalities throughout the Caribbean and western Atlantic, the two species are now re-expanding their ranges northward along the Florida Peninsula and into the northern Gulf of Mexico, coincident with increasing sea temperatures. In the face of continued global warming, the northernmost limit of this range expansion will ultimately be determined by a combination of temperature and other physical constraints.

Reaser, JK, Pomerance, R and Thomas, PO. (2000) Coral bleaching and global climate change: Scientific findings and policy recommendations. Conservation Biology 14, 1500-1511. In 1998, tropical sea surface temperatures were the highest on record, topping off a 50-year trend for some tropical oceans. In the same year, coral reefs around the world suffered the most extensive and severe bleaching (loss of symbiotic algae) and subsequent mortality on record, These events may not be attributable to local stressors or natural variability alone but were likely induced by an underlying global phenomenon. It is probable that anthropogenic global warming has contributed to the extensive coral bleaching that has occurred simultaneously throughout the reef regions of the world. The geographic extent, increasing frequency, and regional severity of mass bleaching events are an apparent result of a steadily rising baseline of marine temperatures, combined with regionally specific El Niño and La Niña events. The repercussions of the 1998 mass bleaching and mortality events will be far-reaching. Human populations dependent on reef services face losses of marine biodiversity, fisheries, and shoreline protection. Coral bleaching events may become more frequent and severe as the climate continues to warm, exposing coral reefs to an increasingly hostile environment. This global threat to corals compounds the effects of more localized anthropogenic factors that already place reefs at risk. Significant attention needs to be given to the monitoring of coral reef ecosystems, research on the projected and realized effects of global climate change, and measures to curtail greenhouse gas emissions. Even those reefs with well-enforced legal protection as marine sanctuaries, or those managed for sustainable use, are threatened by global climate change. Reynaud, S, LeClercq, N, Romaine-Lious, S, Ferrier-Pagès, C, Jaubert, J and Gattuso, J-P. (2003) Interacting effects of CO2 partial pressure and temperature on photosynthesis and calcification in a scleractinian coral. Global Change Biology 9, 1660–1668. We show here that CO2 partial pressure (pCO2) and temperature significantly interact on coral physiology. The effects of increased pCO2 and temperature on photosynthesis, respiration and calcification rates were investigated in the scleractinian coral Stylophora pistillata. Cuttings were exposed to temperatures of 25 1C or 281C and to p CO2 values of ca. 460 or 760 latm for 5 weeks. The contents of chlorophyll c2 and protein remained constant throughout the experiment, while the chlorophyll a content was significantly affected by temperature, and was higher under the ‘high-temperature–high-pCO2’ condition. The cell-specific density was higher at ‘high pCO2’ than at ‘normal pCO2’ (1.7 vs. 1.4). The net photosynthesis normalized per unit protein was affected by both temperature and pCO2, whereas respiration was not affected by the treatments. Calcification decreased by 50% when temperature and pCO2 were both elevated. Calcification under normal temperature did not change in response to an increased pCO2. This is not in agreement with numerous published papers that describe a negative relationship between marine calcification and CO2. The confounding effect of temperature has the potential to explain a large portion of the variability of the relationship between calcification and pCO2 reported in the literature, and warrants a reevaluation of the projected decrease of marine calcification by the year 2100.

Richier, S, Furla, P, Plantivaux, A, Merle, PL and Allemand, D. (2005) Symbiosis-induced adaptation to oxidative stress. The Journal of Experimental Biology 208, 277-285. Cnidarians in symbiosis with photosynthetic protists must withstand daily hyperoxic/anoxic transitions within their host cells. Comparative studies between symbiotic (Anemonia viridis) and non-symbiotic (Actinia schmidti) sea anemones show striking differences in their response to oxidative stress. First, the basal expression of SOD is very different. Symbiotic animal cells have a higher isoform diversity (number and classes) and a higher activity than the non-symbiotic cells. Second, the symbiotic animal cells of A. viridis also maintain unaltered basal values for cellular damage when exposed to experimental hyperoxia (100% O(2)) or to experimental thermal stress (elevated temperature +7 degrees C above ambient). Under such conditions, A. schmidti modifies its SOD activity significantly. Electrophoretic patterns diversify, global activities diminish and cell damage biomarkers increase. These data suggest symbiotic cells adapt to stress while non-symbiotic cells remain acutely sensitive. In addition to being toxic, high O(2) partial pressure (P(O(2))) may also constitute a preconditioning step for symbiotic animal cells, leading to an adaptation to the hyperoxic condition and, thus, to oxidative stress. Furthermore, in aposymbiotic animal cells of A. viridis, repression of some animal SOD isoforms is observed. Meanwhile, in cultured symbionts, new activity bands are induced, suggesting that the host might protect its zooxanthellae in hospite. Similar results have been observed in other symbiotic organisms, such as the sea anemone Aiptasia pulchella and the scleractinian coral Stylophora pistillata. Molecular or physical interactions between the two symbiotic partners may explain such variations in SOD activity and might confer oxidative stress tolerance to the animal host. Ridgway, T. (2005) Allozyme electrophoresis still represents a powerful technique in the management of coral reefs. Biodiversity and Conservation 14, 135–149. Understanding genetic variability and gene flow between populations of scleractinian corals separated by one to several hundred kilometers is crucially important as we head into a century of climate change in which an understanding of the connectivity of populations is a critically important question in management. Genetic methods that directly use molecular variance in the DNA should offer greater precision in detecting differences among individuals and populations than the more traditional allozyme electrophoresis. However, this paper highlights the point that the limited number of DNA markers that have been identified for scleractinian coral genetic studies do not necessarily offer greater precision than that offered by allozymes. In fact, at present allozyme electrophoresis yields greater information than the eight different DNA markers used in this study. Given the relative ease of use of allozymes and the wealth of comparable data sets from numerous previously published studies, allozyme electrophoresis should not be dismissed for population structure and connectivity studies on coral reefs. While continued effort should be placed into searching for new DNA markers, until a more sensitive DNA marker becomes available for scleractinian corals, allozyme electrophoresis remains a powerful and relevant technique for understanding the connectivity of coral population studies.

Riegl, B. (2002) Effects of the 1996 and 1998 positive sea-surface temperature anomalies on corals, coral diseases and fish in the Arabian Gulf (Dubai, UAE). Marine Biology 140, 29-40. Two positive sea-surface temperature anomalies occurred in the Arabian Gulf in short sequence. Between May and August 1996 and 1998, sea-surface temperatures in the southern Arabian Gulf were elevated by 2°C above average. The consequences for coral fauna, coral diseases and coral regeneration were studied in Dubai (United Arab Emirates) between Jebel Ali and Ras Hasyan. In 1996, coral death was widespread, affecting primarily the genus Acropora. In Acropora-dominated areas, live coral cover was reduced from 90% to about 26% in 1996, while in 1998 only a reduction from 26% to 22% of the remaining coral cover occurred. In the study area, all six Acropora species suffered total mortality in 1996, thus the coral fauna was reduced from 34 species to 27. The nearest areas with surviving Acropora were 30 km to the east (Deira) and 20 km to the west (Al Jazira). Massive coral species suffered negligible mortality, and slowly increased in space cover. The Acropora overkill turned 7.9 km2 (19.7% of total coral-covered area) of previously lush coral gardens into a dead framework that was increasingly bioeroded. Acropora recruitment only started in 1998, average recruit size in 1999 was 7±3 cm, and recruits were rare. Prior to the mass mortality event, coral diseases were common and seasonal (14±5% of corals, mainly Acropora, affected in summer, in winter 7±6%, mainly massives), after the mortality event seasonality was lost and infection remained below winter levels (6±5%, only massives infected). In fish, overall species richness decreased from 95 to 64 species in point counts, but frequency only decreased in one species (Pseudochromis persicus). Guild structure changed inasmuch as herbivores and planktivores increased, and invertivores decreased, although differences were not statistically significant. The most abundant family, both prior to and after the coral mass mortality, was Lutjanidae. It appears that even though much of the coral was dead, the maintenance of structural complexity allowed the fish assemblage to avoid a similar catastrophic change to that experienced by the coral assemblage. Riegl, B. (2003) Climate change and coral reefs: Different effects in two high-latitude areas (Arabian Gulf, South Africa). Coral Reefs 22, 433-446. The findings in this paper show that Arabian Gulf (Abu Dhabi, Dubai, Sharjah) corals have already been measurably affected by climate change and further negative impacts are expected. Corals in South Africa have been only weakly impacted and are expected to persist in this likely refuge. The Arabian Gulf has recently experienced high-frequency recurrences of temperature-related bleaching (1996, 1998, 2002). First evidence may suggest that bleaching patterns in corals changed due to phenotypic adaptation after two strong bleaching events in rapid succession, because Acropora, which during the 1996 and 1998 events always bleached first and suffered heaviest mortality, bleached less than all other corals in 2002 at Sir Abu Nuair and recovered at Jebel Ali and Ras Hasyan. In South Africa, reef corals largely escaped the mass mortalities observed across the tropics in the late 1990s, although bleaching has also increased since 1999. These reefs are protected by local small-scale upwelling events in summer that, if they occur at the right time, keep temperatures below bleaching levels. Both areas, the Arabian Gulf and South Africa, have rich coral faunas but little to no recent reefframework production. It is possible

that many reefs worldwide may have similar dynamics in the future, if the changed climate (recurrence of temperature anomalies, changes in aragonite saturation state, etc.) suppresses sustained reef building at least temporarily. Global climate models predict the possibility of significant environmental changes, including increases in atmospheric temperature, sea-surface temperature (SST), and sea level. Monsoon and El Niño Southeastern Oscillation (ENSO) patterns might change, but climate models are not conclusive. Sea-level rise by up to 0.88 m is expected to be a problem in some low-lying areas, like the southern Arabian Gulf. Ocean aragonite saturation state is predicted to fall throughout the ocean but may not change reef dynamics in the two study areas. Riegl, B and Piller, WE. (2003) Possible refugia for reefs in times of environmental stress. International Journal of Earth Sciences 92, 520-531. This paper investigates the refuge potential of (1) upwelling areas, (2) coral areas at medium depth, and (3) offshore bank and island reefs in a scenario of increased global warming, and thus increased sea surface temperature (SST) and increased solar UV radiation. (1) Observations on coral health and water temperature in the subtropical Atlantic (Eleuthera and Cat Island, Bahamas) and Indian Ocean (Sodwana Bay, South Africa) suggest a link between cool water delivered by upwelling and coral health. After the 1998 bleaching event, caused by strong SST anomalies, coral health and recovery from the previous year’s bleaching was significantly better on the narrow southern Cat Island shelf (70% of corals healthy) where the presence of cold water was observed, which was attributed to small-scale upwelling, than on the wide northern Eleuthera shelf (44% of corals healthy), where downwelling of hot bank waters was believed to have damaged corals. In South Africa, regular, short-term upwelling events in five summers reduced SST to well below bleaching level. (2) In the northern Red Sea (Safaga Bay) and in South Africa (Sodwana Bay), wide areas with either coral frameworks or non-framework communities exist. Calculations show that if the top 10 m (20 m) of the ocean became inhospitable to corals, still 50.4% (17.5%) of the coral area would remain intact in the Red Sea and 99% (40%) in South Africa. (3) Offshore bank and island reefs investigated in the Turks, Caicos, and Mouchoir Banks and Grand and Little Cayman showed high rates of mortality and coral diseases. The most remote sites (Mouchoir Bank) were not the healthiest. Refuge areas appear to exist in (1) and (2), but in (3) only if vigorous water-circulation is encountered. Rinkevich, B. (2000) Steps towards the evaluation of coral reef restoration by using small branch fragments. Marine Biology 136, 807-812. “Gardening'' of denuded coral reef habitats is a novel restoration approach in which sexual and asexual recruits are used. The present study aimed at the evaluation of the potentiality for restoration use of different types of small fragments subcloned from the Red Sea coral species Stylophora pistillata. In situ short-term (24 h, 45Ca method) and long-term (1 year, alizarin Red S vital staining) experiments revealed high variation (up to 70%) in growth rates between up-growing branches of a specific genet, and that tip ratios in dichotomous branches (n = 880) differ significantly between newly formed and older branches, further emphasizing the within-colony genetic background for spatial configuration. Small, isolated branches (<4 cm) revealed high survivorship (up to 90%, 1 year) and up to 20-30% (1 year, single- vs.

dichotomous-tip branches, respectively) growth, showing that small-sized branches are suitable for restoration purposes. Results differed significantly between genets. Total length added for dichotomous-tip branches was in general at least twice that recorded for single tips of a specific genet. Restoration protocols may be applied either by sacrificing whole large colonies via pruning high numbers of small fragments or, by pruning only a few small branches from each one of many genets. An in situ “nursery period'' of approximately 8 years is predicted for S. pistillata small fragments. Roberts, CM. (1997) Connectivity and management of Caribbean coral reefs. Science 278, 1454-1457. Surface current patterns were used to map dispersal routes of pelagic larvae from 18 coral reef sites in the Caribbean. The sites varied, both as sources and recipients of larvae, by an order of magnitude. It is likely that sites supplied copiously from “upstream” reef areas will be more resilient to recruitment overfishing, less susceptible to species loss, and less reliant on local management than places with little upstream reef. The mapping of connectivity patterns will enable the identification of beneficial management partnerships among nations and the design of networks of interdependent reserves. Roberts, CM and Hawkins, JP. (2000) Fully-protected marine reserves: A guide. WWF: Washington DC (USA), 131 pp. Fully-protected marine reserves are areas of the sea completely protected from fishing and other extractive or harmful human uses. Since the first fully-protected reserves were established, more than two decades ago, they have stimulated a wealth of research and intense interest. Recent scientific evidence indicates that reserves are not only powerful tools for conservation, but can also provide much needed support for fisheries. There is an urgent need for more reserves in order to address the developing crisis in the oceans. Worldwide, fisheries are in trouble, and habitats and species are being lost at an alarming rate. However, decision makers need good scientific information on how to make reserves work successfully. Questions such as ‘how do reserves function?’, ‘how many should we have?’, and ‘where should we put them?’ are challenging the minds of scientists, conservationists and managers everywhere. The case for marine reserve establishment gets stronger with every new study published and scientists are making good headway in developing a detailed theoretical basis for fully protected reserves, supported by good quality data. People responsible for establishing marine reserves are rarely scientists. Few of those who lobby hardest will have a doctorate in fisheries biology or ecology, nor will the people who decide whether or not to implement protection. People who fish and whose livelihoods will be directly affected by reserves, are educated by the sea itself. Yet all of them, be they fishers, conservationists or government ministers, need clear answers to basic questions and concerns about reserves. For any nonspecialist, whatever their level of education, this can be problematic. Scientific papers are difficult to read and can be hard to acquire. Scientific research can also take years from completion to publication as it grinds through peer review, then joins the queue for a journal slot. The most recent research, while much talked about among scientists, is thus generally inaccessible to those who need it most. The aim of this information pack is to summarize the

scientific case for fully-protected reserves in a way that is easily understood by everyone. Our objective in producing it is to speed up the process of translating scientific research into action. The pack is particularly aimed towards people who need information to inform and persuade others of the benefits of reserves. They include, for example, those working to set up community-based management of marine resources, park or fishery managers, and policy makers. Since people who will be affected by reserves must be willing to place their faith, and possibly risk their livelihoods, on conclusions drawn by scientists, they should be in no doubt about why they are doing so. In addition to explaining the theory behind fully-protected reserves this book is also intended as a practical guide. The main text of the book provides much of the background to reserves. This is supplemented by a series of case studies showing some of the most interesting reserves from around the world. Each of them highlights key findings and identifies lessons from those cases. Two other features of this pack are designed to help people trying to establish reserves in the field. A collection of 30 slides and accompanying descriptions showcase marine reserves and their benefits. They can be used as the basis for a presentation on reserves, either in their entirety, or split into shorter presentations highlighting particular uses of reserves for different audiences. A series of 12 overhead transparencies, with accompanying text, can also be used as the basis for one or more tailored presentations, and could be used to accompany a slide-based presentation. As new information is constantly emerging, we intend this book to be a living text that we update regularly, adding new case studies and sections. These updates can be downloaded from the website that will accompany the book (which can be accessed via a link from www.panda.org/endangeredseas/). We welcome your suggestions for new sections, or case studies that you would like to see covered. You can email them to us at cr10@york.ac.uk. Roberts, CM, McClean, CJ, Veron, JEN, Hawkins, JP, Allen, GR, McAllister, DE, Mittermeier, CG, Schueler, FW, Spalding, M, Wells, F, Vynne, C and Werner, TB. (2002) Marine biodiversity hotspots and conservation priorities for tropical reefs. Science 295, 1280-1284. Coral reefs are the most biologically diverse of shallow water marine ecosystems but are being degraded worldwide by human activities and climate warming. Analyses of the geographic ranges of 3235 species of reef fish, corals, snails, and lobsters revealed that between 7.2% and 53.6% of each taxon have highly restricted ranges, rendering them vulnerable to extinction. Restricted-range species are clustered into centers of endemism, like those described for terrestrial taxa. The 10 richest centers of endemism cover 15.8% of the world’s coral reefs (0.012% of the oceans) but include between 44.8 and 54.2% of the restricted-range species. Many occur in regions where reefs are being severely affected by people, potentially leading to numerous extinctions. Threatened centers of endemism are major biodiversity hotspots, and conservation efforts targeted toward them could help avert the loss of tropical reef biodiversity.

Robison, JD and Warner, ME. (2006) Differential impacts of photoacclimation and thermal stress on the photobiology of four different phylotypes of Symbiodinium (Pyrrhophyta). Journal of Phycology 42, 568-579. The capacity for photoacclimation to light at 100 or 600 lmol photons ·m-2 · s-1 and the subsequent response to thermal stress was examined in four genetically distinct cultures of symbiotic dinoflagellates in the genus Symbiodinium with the ITS2 designations A1, A1.1, B1, and F2. While all algal types showed typical signs of photoacclimation to high light via a reduction in chl a, there was a differential response in cellular growth, photosystem II (PSII) activity, and the chl a-specific absorption coefficient between cultures. When maintained at 321 C for up to 10 days, significant variation in the susceptibility to thermal stress was observed in the rate of loss in PSII activity and electron transport, PSII reaction center degradation, and cellular growth. The order of thermal tolerance did not change between the two light levels. However, as expected, loss in photosynthetic function was exacerbated in the thermally sensitive phylotypes (B1 and A1.1) when acclimated to the higher light intensity. There was no consistent relationship between thermal tolerance and changes in light energy dissipation via non-photochemical pathways. Phylotypes F2 and A1 showed a high degree of thermal tolerance, yet the cellular responses to light and temperature were markedly different between these algae. The F2 isolate showed the greatest capacity for photoacclimation and growth at high light and temperature, while the A1 isolate appeared to adjust to thermal stress by a slight decline in PSII activity and a significant decline in growth, possibly at the expense of increased photosystem and cellular repair rates. Rodolfo-Metalpa, R, Richard, C, Allemand, D, Bianchi, CN, Morri, C, Ferrier-Pagès, C. (2006) Response of zooxanthellae in symbiosis with the Mediterranean corals Cladocora caespitosa and Oculina patagonica to elevated temperatures. Marine Biology 150, 45-55. Scleractinian symbiotic corals living in the Ligurian Sea (NW Mediterranean Sea) have experienced warm summers during the last decade, with temperatures rapidly increasing, within a few days, to 3–4°C above the mean value of 24°C. The effect of elevated temperatures on the photosynthetic efficiency of zooxanthellae in symbiosis with temperate corals has not been well investigated. In this study, the corals, Cladocora caespitosa and Oculina patagonica were collected in the Ligurian Sea (44°N, 9°E), maintained during 2 weeks at the mean summer temperature of 24°C and then exposed during 48 h to temperatures of 24 (control), 27, 29 and 32°C. Chlorophyll (chl) fluorescence parameters [Fv/Fm, electron transport rate (ETR), non-photochemical quenching (NPQ)] were measured using pulse amplitude modulated (PAM) fluorimetry before, during the thermal increase, and after 1 and 7 days of recovery (corals maintained at 24°C). Zooxanthellae showed a broad tolerance to temperature increase, since their density remained unchanged and there was no significant reduction in their maximum quantum yield (Fv/Fm) or ETR up to 29°C. This temperature corresponded to a 5°C increase compared to the mean summer temperature (24°C) in the Ligurian Sea. At 32°C, there was a significant decrease in chl contents for both corals. This decrease was due to a reduction in the chl/zooxanthellae content. For C. caespitosa, there was also a decrease in ETRmax, not associated with a change in Fv/Fm or in the non-photochemical quenching (NPQ); for O. patagonica, both ETRmax and Fv/Fm significantly decreased, and NPQmax

showed a significant increase. Damages to the photosystem II appeared to be reversible in both corals, since Fv/Fm values returned to normal after 1 day at 24°C. Zooxanthellae in symbiosis with the Mediterranean corals investigated can therefore be considered as resistant to short-term increases in temperature, even well above the maximum temperatures experienced by these corals in summer. Rosenberg, E and Loya, Y. (2004) Coral health and disease. Springer Verlag: Berline (Germany), 488 pp. Coral reefs are the most spectacular and diverse ecosystems in the marine environment. Over the last decades, however, dramatic declines of coral reef communities have been observed. Corals are endangered due to natural and anthropogenic detrimental factors, such as global warming and environmental pollution. Based on an international meeting on "Coral Health and Disease" in Eilat, Israel in April 2003, the book starts with case studies of reefs, e.g. the Red Sea, Caribbean, Japan, Indian Ocean and the Great Barrier Reef. The second part on microbial ecology and physiology describes the symbiotic relations of corals and microbes, and the microbial role in nutrition or bleaching resistance of corals. Particular coral diseases such as aspergillosis, white pox, black and white band diseases are treated in the third part. Finally, various hypotheses of the mechanisms of coral bleaching, including a projection of the future of coral reefs, are discussed. Rotjan, RD, Dimond, JL, Thornhill, DJ, Leichter, JJ, Helmuth, B, Kemp, DW and Lewis, SM. (2006) Chronic parrotfish grazing impedes coral recovery after bleaching. Coral Reefs 25, 361-368. Coral bleaching, in which corals become visibly pale and typically lose their endosymbiotic zooxanthellae (Symbiodinium spp.), increasingly threatens coral reefs worldwide. While the proximal environmental triggers of bleaching are reasonably well understood, considerably less is known concerning physiological and ecological factors that might exacerbate coral bleaching or delay recovery. We report a bleaching event in Belize during September 2004 in which Montastraea spp. corals that had been previously grazed by corallivorous parrotfishes showed a persistent reduction in symbiont density compared to intact colonies. Additionally, grazed corals exhibited greater diversity in the genetic composition of their symbiont communities, changing from uniform ITS2 type C7 Symbiodinium prior to bleaching to mixed assemblages of Symbiodinium types post-bleaching. These results suggest that chronic predation may exacerbate the influence of environmental stressors and, by altering the coral-zooxanthellae symbiosis, such abiotic-biotic interactions may contribute to spatial variation in bleaching processes. Rotjan, RD and Lewis, SM. (2006) Parrotfish abundance and selective corallivory on a Belizean coral reef. Journal of Experimental Marine Biology and Ecology 335, 292–301. Parrotfish are important members of coral reef communities because they consume macroalgae that would otherwise outcompete reef-building corals for space. However, some Caribbean parrotfish species also feed directly on live corals, and thus have the potential to negatively impact coral fitness and survival. This study investigates selective grazing by parrotfish on particular coral species, differences in

grazing incidence among reef habitats and intraspecific discrimination among colonies of several coral species. We also investigate spatial and temporal patterns of parrotfish species abundance across habitats on the Belize barrier reef, and examine correlations between parrotfish abundance and grazing intensity across reef habitats. We found that members of the Montastraea annularis species complex, major builders of Caribbean reefs, were preferred targets of parrotfish grazing across all reef habitats, while M. cavernosa, Agaricia agaricites, Diploria strigosa, Porites astreoides and Porites porites were not preferred; Siderastrea siderea was preferentially grazed only in the spur and groove habitats. Parrotfish grazing preferences varied across habitats; M. annularis was grazed most often in shallow habitats, whereas M. franksi was consumed more at depth. Although it was not possible to directly observe parrotfish grazing on corals, we did find a positive correlation between Sparisoma aurofrenatum abundance and M. franksi grazing incidence across habitats. Finally, when we compared our results to parrotfish abundances measured by a previous study, we found that Sparisoma viride and Sp. aurofrenatum, two species known to be corallivorous, had increased abundances between 1982 and 2004. In light of escalating threats on Caribbean reef corals, it would be important for future studies to evaluate the impact of parrotfish corallivory on coral survival. Rowan, R, Knowlton, N, Baker, A and Jara, J. (1997) Landscape ecology of algal symbionts creates variation in episodes of coral bleaching. Nature 388, 265-269. Reef-building corals are obligate, mutualistic symbioses of heterotrophic animals and phototrophic dinoflagellates (Symbiodinium spp.). Contrary to the earlier, widely accepted belief that corals harbour only one symbiont, we found that the ecologically dominant Caribbean corals Montastraea annularis and M. faveolata can act as hosts to dynamic, multi-species communities of Symbiodinium. Composition of these communities follows gradients of environmental irradiance, implying that physiological acclimatization is not the only mechanism by which corals cope with environmental heterogeneity. The importance of this diversity was underlined by analysis of a natural episode of coral bleaching. Patterns of bleaching could be explained by the preferential elimination of a symbiont associated with low irradiance from the brightest parts of its distribution. Comparative analyses of symbionts before and after bleaching from the same corals supported this interpretation, and suggested that some corals were protected from bleaching by hosting an additional symbiont that is more tolerant of high irradiance and temperature. This 'natural experiment' suggests that temporal and spatial variability can favour the coexistence of diverse symbionts within a host, despite the potential for destabilizing competition among them. Rowan, R. (2004) Coral bleaching: Thermal adaptation in reef coral symbionts. Nature 430, 742. Many corals bleach as a result of increased seawater temperature, which causes them to lose their vital symbiotic algae (Symbiodinium spp.) - unless these symbioses are able to adapt to global warming, bleaching threatens coral reefs worldwide. Here I show that some corals have adapted to higher temperatures, at least in part, by

hosting specifically adapted Symbiodinium. If other coral species can host these or similar Symbiodinium taxa, they might adapt to warmer habitats relatively easily. Ruiz-Zárate, MA and Arias-González, JE. (2004) Spatial study of juvenile corals in the Northern region of the Mesoamerican Barrier Reef System (MBRS). Coral Reefs 23, 584-594. Juvenile coral abundance and community composition depend on the spatial scale studied. To investigate this, an evaluation was made of juvenile coral density with hierarchical spatial analysis in the northern Mesoamerican Barrier Reef System (MBRS) at ~10 m depth. Study scope included semi-protected and unprotected areas located in this region. A total of 19 juvenile coral taxa were found, including 10 scleractinian species, 8 scleractinian coral genera not identified to species, and 1 Millepora species (Hydrozoa-Milleporidae). In terms of relative abundance, Agaricia spp., Siderastrea spp., and Porites spp. were the main juvenile taxa in the coral community at the surveyed sites, reefs, and regions levels. Greater variance was seen at smaller scales, at site level for taxa richness, and at the transect level for juvenile density, and lower variance was seen at larger scales (reefs and regions). The variance component contribution from each scale likely differed from other studies because of the different factors affecting the community and the different extensions of each scale used in each study. Densities (1–6.4 juvenile corals/m2) and dominant taxa found in this study agree with other studies from the Western Atlantic. Detected variability was explained by different causal agents, such as low grazing rates by herbivorous organisms, turbidity, and/or sediment suffocation and some nearby or distant localized disturbance (human settlement and a hurricane). Salih, A, Larkum, A, Cox, G, Kühl, M and Hoegh-Guldberg, O. (2000) Fluorescent pigments in corals are photoprotective. Nature 408, 850-853. All reef-forming corals depend on the photosynthesis performed by their algal symbiont, and such corals are therefore restricted to the photic zone. The intensity of light in this zone declines over several orders of magnitude - from high and damaging levels at the surface to extreme shade conditions at the lower limit. The ability of corals to tolerate this range implies effective mechanisms for light acclimation and adaptation. Here we show that the fluorescent pigments (FPs) of corals provide a photobiological system for regulating the light environment of coral host tissue. Previous studies have suggested that under low light, FPs may enhance light availability. We now report that in excessive sunlight FPs are photoprotective; they achieve this by dissipating excess energy at wavelengths of low photosynthetic activity, as well as by reflecting of visible and infrared light by FP-containing chromatophores. We also show that FPs enhance the resistance to mass bleaching of corals during periods of heat stress, which has implications for the effect of environmental stress on the diversity of reef-building corals, such as enhanced survival of a broad range of corals allowing maintenance of habitat diversity.

Salm, RV, Clark, JR and Siirila, E. (2000) Marine and Coastal Protected Areas: A guide for planners and managers. IUCN: Gland (Switzerland), 371 pp. This book is addressed mainly to conservation of the natural resources of tropical coasts and seas. To facilitate its use as a sourcebook, the volume is arranged in three parts. Part I introduces Marine Protected Areas (MPAs) as an important approach to managing coastal and marine resources, discussing in its six sections: roles of MPAs, site planning, community involvement, systematic selection of MPAs, strategies and tools for planning and managing MPAs, and the legal basis of MPAs. Part II considers principles and mechanisms for planning and managing protected areas in four different environments: coral reefs, estuaries and lagoons, small islands, and beaches. Emphasis is on technical knowledge about particular habitats and how this knowledge is used to meet management objectives. We recognize that many MPAs are composed of two or more of these environments but the guidance for each can be combined in the MPA plan. Part III presents case histories covering a wide variety of MPA experience around the world to help protected area planners and managers carry out their tasks. The emphasis in each case history is on lessons learned that are of wide application. E-mail addresses of the authors are given to assist the reader in following up on case details. A “Guide to the Cases” is presented to enhance the reader’s search for relevant items. In the selection of materials for this book we have given priority to the types of resource conservation problems faced by MPA planners and managers in subtropical and tropical countries where the need for assistance appears to be the greatest. In Parts I and II most of the material is based on written sources for which the specific references are listed alphabetically with date of publication in Part IV, References. Case Histories are individually referenced. Salm, RV and Coles, SL (eds.) (2001) Coral bleaching and Marine Protected areas. Proceedings of the Workshop on Mitigating Coral Bleaching Impact Through MPA Design, The Nature Conservancy: Honlulu (USA), 118 pp. Those of us who knew the corals reefs of the world 30 years ago or more have seen widespread deterioration in their condition and in the numbers of fishes and mollusks they shelter. Projecting the past 30 years of change ahead another 30 years might be enough to drag us into a vortex of despondency. On the other hand, inspired by our conservation successes, we could rise to the challenge and look for solutions. It makes good sense to prepare for what lies ahead in this new century by looking back at changes in the state of reefs and projecting these trends ahead to anticipate emerging threats. In doing so, we can position ourselves to be ready for them. Bleaching as a response by corals to stress caused by elevated seawater temperature is one of these emerging threats that has already led to the possible extinction of some coral species from reefs in the Panama and Okinawa regions, at opposite ends of the Pacific. Mass coral bleaching of the scale following the 1998 El Niño Southern Oscillation (ENSO) event is viewed by some as an intractable problem and by others as a challenge we need to do "something" about. Patterns of bleaching in the Florida Keys National Marine Sanctuary show that each subsequent event has exceeded former thresholds of resistance and extended bleaching to new reefs. This is a matter of

concern that increases the stakes and warns us that we cannot wait any longer to act. Responding to rising global concern, The Nature Conservancy (TNC) joined with the World Wildlife Fund U.S. (WWF-US) to determine how to shape the "something" into practical actions that could be implemented to mitigate the impacts of bleaching on coral reef biodiversity. We convened a small working group of influential participants with significant experience in relevant coral reef research, monitoring, and marine protected area (MPA) management. Collectively, they embody comprehensive, direct knowledge of all major reef areas worldwide (Asia, Pacific, Indian Ocean, Caribbean, United States, and Australia). The group was united and stimulated by a shared concept that, counter to the prevailing sentiment of doom and gloom, something can be done to address the severe impacts of climate related bleaching on coral reefs, even if sources of climate change are beyond our control within meaningful timeframes. This first workshop was a determined attempt to explore and agree on some potential science-based principals that could guide management of current MPAs to mitigate the impacts of coral bleaching, optimize conditions for reef recovery, and steer the establishment of new MPAs. The intent was to build upon, not change, the many varied and excellent approaches to reef management by adding some complementary activities and approaches that are designed specifically to help managers address coral bleaching through mitigation strategies. Hopefully, the workshop will catalyze a succession of other small, focused meetings aimed at determining how to control or mitigate other unmanageable global threats to marine biodiversity that are likely to emerge in this century. Examples of these include: sea level rise; increased demand for living space, food, materials, recreation, and tourism; habitat fragmentation and isolation; and accelerated introduction of invasive species. The group confirmed a concept first developed by TNC that certain environmental factors favor survival or recovery of corals and other organisms which have bleached, and that policy-makers and resource managers should include reefs containing such factors within MPA networks. Also stressed was the need to assist the recovery of coral reefs that have bleached by protecting them from siltation, pollution, overfishing, and other destructive practices. The group recognized that areas resistant to bleaching are critical to the recovery and survival of coral communities that succumb to bleaching or other stresses. If these resistant sites are not protected, they may be degraded by a host of other factors and lose their effectiveness as sources of larvae to enable the recovery of affected areas. Consequently, management should aim to protect areas of high resistance to bleaching, but protection should also be extended to enhance the recovery of adjacent areas by reducing the influence of other stresses. The group also found value in increasing the probability of reef community recovery from bleaching and other stresses through replication of MPAs—the more MPAs there are, the greater the likelihood that significant components of these will survive to facilitate recovery of affected areas. The group agreed on a list of factors that confer bleaching resistance (coral colonies that don’t bleach or bleach but don’t die) and resilience (reefs where colonies bleach and partially or wholly die, but the coral community recovers). They outlined a general approach for a global program to test and verify that environmental factors are effective in mitigating bleaching impacts. The goal of this global program is to maximize the protection of coral reefs in the face of expected increases in frequency and intensity of climate related bleaching events.

Furthermore, the group agreed to work with MPA practitioners around the world to evaluate the vulnerability of existing MPAs to bleaching and the practicality of implementing the new management approaches. They recommended that additional criteria and principles linked to survivability should be considered alongside the widely accepted ones when selecting and designing new coral reef MPAs, but stressed the need to assist the recovery of bleached reefs by protecting them from other threats and managing them to promote resilience. To derive all essential empirical data to support additional practices in MPA establishment and management could take up to a decade of research and monitoring. In the interim, there is value in using the best available information, both observational and available scientific data, to define MPA selection criteria, design principles, and management guidelines that can be updated and refined as the years go by. The ongoing research and monitoring also will be integral for developing an understanding of how to manage coral reefs in the face of global change. The group agreed to proceed along a two-track approach. The first track includes worldwide assessments, research, and monitoring. These will be designed to provide solid scientific evidence to verify current information regarding the contribution of environmental factors to bleaching resistance and resilience. The reliability of these factors will be tested and applied to generate additional MPA selection criteria and design principles—the two primary components of the second track. Simultaneously, through the second track, draft additional MPA selection criteria and design principles will be prepared and distributed globally for application, verification, and refinement. The IUCN World Commission on Protected Areas (WCPA) will play a key role in this process. The target is to assimilate feedback from the two tracks and present and discuss these additional MPA criteria and principles at the 5th World Parks Congress, to be convened by IUCN WCPA in mid-2003. A global MPA practitioners support network will be established during the Congress to facilitate global application of the new selection criteria and design principles. Just as bleaching is not exclusive to a country or region and recognizes no institutional or national boundaries, neither is exclusion the intention of this two-track program, which would clearly benefit by implementation through a global partnership. TNC, WWF, and the workshop participants expressed firm commitment to build on this successful workshop and to collaborate to take this initiative forward in partnership with any other interested private or public organizations. Salm, RV. (2002) Preparing Marine Protected Areas to survive global change: Additional guidelines to address coral bleaching. The Nature Conservancy: Honolulu (USA), 19 pp. In the face of rampant reef destruction and the staggering impacts of such large-scale unmanageable threats as climate-related thermal bleaching, people are beginning to ask whether there is anything we can do to conserve our coral reefs. The greater skeptics among these even venture to ask “why bother trying to conserve coral reefs when events that are beyond our control will undermine our best efforts?” The answer to this question would certainly be “we probably shouldn’t bother trying if we keep doing what we have always done. However, if we can adapt our approaches to the times, we should never stop trying!” The good news is that for every statistic that describes the corals lost to bleaching, there is the complementary statistic that indicates how much coral survives. This largely unspoken statistic gives us hope; and if we can understand the survival

factors underlying it, we will have the means to adjust our approaches to coral reef conservation so that they build on survival, despite these daunting global trends. This document is about moving beyond hope. It is about looking at what survives a major bleaching event, and about building that into our coral reef conservation programs. It is a piece of a larger initiative that originated one sunset in Komodo National Park in Indonesia and had its formal beginnings at the 9th Coral Reef Symposium in October 2000. There, the concept was first presented for discussion that certain environmental factors seem to help some coral communities resist bleaching and that these could be identified and used to influence to how we select, design and manage marine protected areas. Encouraged by the favorable response this presentation received at the symposium, The Nature Conservancy and World Wildlife Fund (US) convened a small working group in 2001 (Salm and Coles 2001). The participants, who collectively had considerable experience in relevant research and MPA management experience in all the major coral reef areas of the world, confirmed the concept that certain environmental factors favor survival and recovery of bleached corals and other organisms. They agreed on a list of factors that confer bleaching resistance (coral colonies that don’t bleach or bleach but don’t die) and resilience (reefs where colonies bleach and partially or wholly die, but the coral community recovers). Furthermore, the participants recommended that policy-makers and resource managers should include reefs containing these factors within MPA networks. Also stressed was the need to assist the recovery of bleached coral reefs by protecting them from siltation, pollution, overfishing, and other destructive practices. The workshop participants outlined a a two-track approach for a global program to test and verify that environmental factors are effective in mitigating bleaching impacts. The goal of this global program is to maximize the protection of coral reefs in the face of expected increases in frequency and intensity of climate related bleaching events. The first track includes worldwide assessments, research, and monitoring. These will be designed to provide solid scientific evidence to verify current information regarding the contribution of environmental factors to bleaching resistance and resilience. The reliability of these factors will be tested and applied to generate additional MPA selection criteria and design principles—the two primary components of the second track. Simultaneously, through the second track, draft additional MPA selection criteria and design principles will be prepared and distributed globally for application, verification, and refinement. This document represents the first step in implementation of the second track. The target is to assimilate feedback from the two tracks and discuss these additional MPA criteria and principles at the 5th World Parks Congress, to be convened by IUCN World Commission on Protected Areas (WCPA) in mid-2003. A need for and structure of a global MPA practitioners support network to facilitate global application of the new selection criteria and design principles will also be discussed during the Congress. The IUCN WCPA will play a key role in this entire process. Recognizing that areas resistant to bleaching are critical to the recovery and survival of coral communities that succumb to bleaching or other stresses, MPA managers should give these sites adequate protection so that they retain their effectiveness as sources of larvae to enable the recovery of affected areas. There is also value in increasing the probability of reef community recovery from bleaching and other stresses through replication of MPAs—the more MPAs there are, the greater the

likelihood that significant components of these will survive to facilitate recovery of affected areas. Salm, RV, Done, T and Tupper, M. (2003) Designing MPAs to survive coral bleaching: What can we do? US Coral Reef Taskforce Coral Reefs, Climate and Coral Bleaching Workshop, 9 pp. The increased frequency and severity of mass coral bleaching over the past decade has increased stresses on coral reefs and raised concerns about appropriate response strategies. This climate related pressure might represent the single greatest short to medium-term threat to coral reefs worldwide. The threat is to both the coral communities per se, and their productivity as essential fish habitat. However, there is a growing body of observational and empirical evidence to suggest that certain coral communities can resist bleaching or, if impacted, recover quickly from it. This paper describes factors that appear to confer resistance and resilience to bleaching and suggests actions that managers may take to build resilience in the face of change into coral reef conservation programs. It advocates the development of a ‘toolkit’ to aid researchers and local MPA practitioners through the complex process of determining and implementing effective local actions. The contents of the toolkit – created and applied collaboratively by scientists with local practitioners – offer assistance in the recognition of key geographic and environmental settings likely to be important in MPA design and management. Salm, RV and West, JM. (2003) Enhancing reef survival in a changing climate: Additional Marine Protected Area guidelines to address coral bleaching. The Nature Conservancy: Honolulu (USA), 21 pp. Large-scale coral bleaching events have increased in intensity, frequency, and geographic distribution in the last two decades (Wilkinson 1998, 2000). The 1998 El Niño Southern Oscillation (ENSO) event and 1999 La Niña caused mass coral bleaching of unprecedented proportions worldwide and near complete loss of live coral at some sites (Goreau et al. 2000). We now recognize that climate-related bleaching events pose a serious global threat to coral reefs (Goreau et al. 2000, Westmacott et al. 2000, Salm et al. 2001); therefore, management and planning guidelines must be expanded to address the challenge. Because ENSO events are expected to occur with greater frequency and intensity over the coming decades, there is an urgent need for practical response measures to mitigate negative impacts on coral communities at sufficiently large scales to be effective (Salm et al. 2001). Despite the widespread bleaching-associated mortality of coral reef organisms, particularly that following the 1998 ENSO, it is rare for living corals to be completely eliminated from a section of reef. Even in the most severe cases, some coral communities appear to be more resistant or more resilient. A recent analysis of bleaching reports (Wilkinson 2000) indicates that there is a wide variability in bleaching intensity, species affected, depth, and geographic distribution, and how much mortality a bleaching event causes. After widespread mortalities from bleaching events have occurred, reefs also differ in their ability to regenerate themselves through growth and expansion of existing corals or recruitment of new corals.

Patterns of bleaching-related mortality and recovery provide insights into the environmental and biological factors that may influence such differential responses to bleaching among various coral reef communities. Based on evidence from the literature and systematically-compiled observations from researchers in the field, we have identified factors that correlate with resistance and resilience to coral bleaching, including a variety of factors that: 1) reduce temperature stress (e.g., upwelling zones); 2) enhance water movement and flush harmful toxins (e.g., narrow channels with strong currents); 3) decrease light stress (e.g., high island shading); 4) correlate with physiological tolerance (e.g., regularly stressful environments where corals are already adapted to stress); and 5) provide physical or biological enhancement of recovery potential (e.g., low abundance of bioeroders and disease) (Salm and Coles 2001, West 2001 a, b, West and Salm 2003). A full list of these factors can be viewed in Appendix A. This information has important implications for identifying reef areas that are likely to be most robust in the face of continuing climate change and for determining priority areas for strict protection from direct anthropogenic impacts. Besides selecting MPAs for the usual range of accepted criteria and managing them as we usually do to control specific threats, we can also introduce additional criteria and design principles focused on reef "survivability" (resistance and resilience). Building survivability into MPA design and management is a new concept. Survivability has never been explicitly defined as a criterion for MPA selection, nor has it been addressed in the principles for MPA design or factored into large scale ecoregional planning. Yet the concept of survivability as described in this report indicates that there are positive actions that we can take to counter the potentially devastating impacts of climate-related bleaching on coral reefs. At this point, we would like to underscore that although existing MPA selection criteria and design principles do not specifically address mass coral bleaching, they retain definite value. They are critical for: 1) defining conservation objectives and targets; 2) identifying threats to these objectives (including sources or root causes of the threats); and 3) determining management strategies to address the threats. These criteria and principles provide the focus for coral reef conservation planning and – along with various approaches, such as community participation, co-management, and the like – provide the core strategies for effective MPA establishment (Salm et al. 2000). With global-scale threats now added to the equation, we need to build onto our existing principles new approaches that complement our usual range of management actions. We must ensure that the areas we conserve are not only protected from the more immediate anthropogenic impacts that we can directly manage, but also are located and managed in a way that boosts their resilience to global-scale impacts that can not be directly managed. This document builds on existing MPA selection criteria, design principles, and management guidelines (e.g., Salm et al. 2000) to present some draft additional guidelines aimed at enhancing the survival and recovery prospects of coral communities affected by ENSO-related bleaching. These additional criteria, principles, and guidelines are derived from concepts first presented at the International Coral Reef Symposium in 2000 (Salm et al. 2001) and expanded in a

subsequent workshop (Salm and Coles 2001, West 2001a,b) and scientific review (West and Salm 2003). Salm, RV, Done, T and McLeod, E. (in press) Marine Protected Area (MPA) planning in a changing climate. In Coral reefs and climate change – Scientific and management implications, American Geophysical Union. The increased frequency and severity of mass coral bleaching due to sea temperature rise has increased stresses on coral reefs and raised concerns about appropriate response strategies (Hughes et al. 2003). Marine Protected Areas (MPAs) have been identified as one of the most effective tools for conserving reefs and related marine systems (Lubchenco et al. 2003; Palumbi 2003). However, protected area managers must incorporate climate change as well as increasing human pressures into their conservation strategies, or MPAs may not be able to safeguard biodiversity effectively. Networks of MPAs, including no-take areas totaling 30-50% of available reef area (Hoegh-Guldberg and Hoegh-Guldberg 2004), have been identified as a critical way to protect coral reefs from human stresses. Implementation of no-take areas has been shown to increase fish biomass and variability both inside and outside the boundaries of those areas (Halpern 2003). Reef fish communities that contain adequate numbers of all key functional groups are believed to contribute to the resilience of coral reef ecosystems – that is, its capacity to recover to a coral-dominated habitat following disturbances. The ideal no-take zone would thus be one that has both inherent resistance to bleaching and all the necessary precursors for resilience, including good on-site conditions and a reliable supply of propagules of all key functional groups. This paper describes environmental factors that appear to confer resistance and resilience to bleaching and suggests actions that MPA managers may take to build resilience in the face of change into coral reef conservation programs. Sammarco, PW, Winter, A and Stewart, JC. (2006) Coefficient of variation of sea surface temperature (SST) as an indicator of coral bleaching. Marine Biology 149, 1337–1344. Coral bleaching has become a major problem on reefs around the world in recent decades. It is believed that mean temperature alone is the primary force driving this ecological phenomenon. We propose that variance in temperature in the short term is just as important as the mean. Thirty years of daily sea surface temperature (SST) data have been collected by the University of Puerto Rico at Mayaguez Marine Laboratory in La Parguera, PR. These data were collated and analyzed initially (by Amos Winter) for their relationship to coral bleaching in this area. We found that the data fell into three categories: high mean temperatures associated with severe bleaching, cooler mean temperatures associated with no bleaching, and years of high SSTs but with no coral bleaching. Here, we examined the relationship between mean temperature during those months in which bleaching occurred, temperature variance (as measured by standard deviation), and coefficient of variation (CV; i.e., SD standardized by the mean). We also derived a critical threshold temperature and level of resolution in time for calculating these statistics to clearly describe the circumstances of bleaching versus non-bleaching events, particularly at marginal bleaching temperatures. These characteristics were compared for the four warmest months of the year (July–October) for four warm bleaching years (1969, 1987, 1990,

and 1995), four cool non-bleaching years (1984, 1985, 1986, and 1988), and two warm nonbleaching years (1994 and 2000). No relationship was found between the mean SST and SD in terms of predicting bleaching. The two primary statistics which, in concert, did indicate bleaching, however, were the shortterm, biweekly mean temperature and its the associated CV. Bleaching occurs in association with both high temperatures and a high CV. The CV becomes a critical determinant of bleaching only when temperatures are ~29.1–29.8°C. The warm, non-bleaching years were generally characterized by a CV of < 1.9 and a temperature range between 28.5 and 29.9°C. We conclude that increased mean SSTs alone are not sufficient to induce coral bleaching; a high variance in SST at marginal, lower bleaching temperatures can induce bleaching, and likewise, a low variance of such will not induce bleaching. This variance is most clearly described by the CV. Samways, MJ. (2005) Breakdown of butterflyfish (Chaetodontidae) territories associated with the onset of a mass coral bleaching event. Aquatic Conservation: Marine and Freshwater Ecosystems 15, S101-S107. 1. Obligate corallivorous butterflyfishes (Chaetodontidae) have been suggested as indicators of the health of coral reefs. 2. There has been a call to relate specific changes in butterflyfish ecology and behaviour to identified stressors. 3. For the first time it is reported here that there is a breakdown of the normal rigid territorial butterflyfish behaviour at the onset of an intense, large-scale coral bleaching event in the Seychelles, western Indian Ocean. 4. Behaviour changes from fixed territory holding, and a mosaic of intra- and interspecific territories, to one of mass excursions by the whole obligate corallivore assemblage. 5. The two main species studied here, Chaetodon trifascialis and C. trifasciatus, fed almost exclusively on Acropora spp. In the western Indian Ocean these species of coral are among the first to die during a bleaching event. 6. It is speculated that the early excursion behavioural response by these butterflyfishes is because their prime food resource is the first to perish. 7. It is recommended that baseline data be gathered on corallivorous butterflyfish territory size and rate of excursion on healthy, unstressed reefs. 8. Deviation from these baseline results, along with early bleaching corals such as Montipora spp. and branching Acropora spp., is highly likely to indicate the coming of a bleaching event. Santavy, DL, Summers, JK, Engle, VD and Harwell, LC. (2005) The condition of coral reefs in South Florida (2000) using coral disease and bleaching as indicators. Environmental Monitoring and Assessment 100, 129-152. The destruction of coral reef habitats has occurred at unprecedented levels during the last three decades. Coral disease and bleaching in the Caribbean and South Florida have caused extensive coral mortality with limited recovery, often coral reefs are being replaced with turf algae. Acroporids were once dominant corals and have diminished to the state where they are being considered as endangered species. Our survey assessed the condition of reef corals throughout South Florida. A probability-based design produced unbiased estimates of the spatial extent of ecological condition, measured as the absence or presence and frequency or prevalence of

coral diseases and bleaching intensity over large geographic regions. This approach allowed us to calculate a quantifiable level of uncertainty. Coral condition was estimated for 4100 hectares (ha) (or 41.0 km2) of coral reefs in South Florida, including reefs in the Florida Keys National Marine Sanctuary (FKNMS), New Grounds, Dry Tortugas National Park (DTNP), and Biscayne National Park (BNP). The absence or presence of coral disease, ‘causal’ coral bleaching, partial bleaching and coral paling were not good indicators of overall coral condition. It was more useful to report the prevalence of anomalies that indicated a compromised condition at both the population and community levels. For example, 79% of the area in South Florida had less than 6% of the coral colonies diseased, whereas only 2.2% (97.15 ha) of the sampled area had a maximum prevalence of 13% diseased coral colonies at any single location. The usefulness of ‘causal bleaching’ might be more important when considering the prevalence of each of the three different states at a single location. For example, paling was observed over the entire area, whereas bleaching and partial bleaching occurred at 19 and 41% of the area, respectively. An index for coral reef condition might integrate the prevalence and species affected by each bleaching state at individual locations. By establishing these baselines, future surveys can examine changes and trends in the spatial distribution of coral conditions in South Florida and able to score the reefs as to their health status. Schlöder, C and D’Croz, L. (2004) Responses of massive and branching coral species to the combined effects of water temperature and nitrate enrichment. Journal of Experimental Marine Biology and Ecology. The branching coral species Pocillopora damicornis (Linnaeus) and the massive coral species Porites lobata Dana were exposed for 30 days to different temperatures and nitrate concentrations to study the response of the coral–zooxanthella symbiosis. Results suggest that the effect of nitrate enrichment on the polyp–zooxanthella symbiosis varies according to the coral morphology. After the experimental period only 30% of P. damicornis colonies remained healthy, in contrast to 90% of P. lobata. The branching P. damicornis was significantly affected by the addition of nitrate, whereas P. lobata was significantly influenced by water temperature. The two species showed enhanced zooxanthella volume, and chlorophyll contents per cell under high nitrate concentrations. The reduced zooxanthellae density in both species indicated a detrimental influence of the interaction of high nitrate and high temperature. Tissue soluble proteins in P. lobata were significantly reduced by elevated temperature. Results showed that tissue soluble proteins and chlorophylls in P. lobata were from two- to three-fold higher than in P. damicornis. The number of zooxanthellae in P. lobata was double that of P. damicornis. Therefore, we suggest that the slow-growing species P. lobata is better able to cope with changing environmental conditions than the fast-growing coral P. damicornis. Schuhmacher, H, Loch, K, Loch, W and See, WR. (2005) The aftermath of coral bleaching on a Maldivian reef – A quantitative study. Facies. Since the bleaching event of 1998, the development of the reef flat and upper reef slope on aMaldivian reef (the Komandoo house reef; Komandoo Island, Lhaviyani atoll or Fadiffolu atoll) is under detailed observation. We quantitatively recorded specific losses, re-colonisation by coral larvae on transects on the reef flat and on dead Acropora tables at the reef slope and regeneration of partly damaged large

Porites and Diploastrea—colonies over the period from 1999 to 2004. The detrimental effects on the reef structure by bioerosion and hydrodynamics, as well as the overall status of the reef community were qualitatively assessed. Recruitment soon after the bleaching was more pronounced than in the following years, Pavona varians being a main constituent. The temporal re-colonisation pattern points at an emergency spawning of local Scleractinia just prior to the bleaching, whereas a sharp decrease of young settlers in 2001 and 2002 confirms a reduction of fertile colonies. The dominant species in the coral community shifted from acroporids and pocilloporids to agariciids. The skeletal deposition of recovering Diploastrea heliopora was equivalent to that before the bleaching, but much less than that of neighbouring Porites lobata colonies. The slow and scattered formation of new reef substance, which would structurally strengthen the reef, is however outweighed by the collapse of dead protruding and spacious colonies (e.g., Acropora tables). Six years after the bleaching, the formerly three-dimensional structure of the reef flat and upper reef slope presents as a levelled field of rubble, only partly consolidated by incrusting corals. Considering the recurrence of bleaching events (1987, 1998) and the results of the present study, one may assume a cascading deterioration of the status of the reef for the future. Schuttenberg, H (ed.) (2001) Coral bleaching: Causes, consequences and responses. Selected papers presented at the 9th International Coral Reef Symposium on ‘Coral bleaching: Assessing and linking ecological and socio-economic impacts, future trends and mitigation planning’ in Bali (Indonesia): Coastal Resources Center, University of Rhode Island (USA), 102 pp. Coral reefs have long fascinated human beings with their myriad of life forms and colors. Coral reefs are among the most biologically diverse ecosystems on the earth, but are degrading at an alarming rate in all tropical oceans. Throughout the Indo-Pacific, only a small fraction of reefs are considered to still be in excellent condition. Many reefs are seriously degraded, particularly in the Philippines and Indonesia. In 1998, the negative impacts of destructive fishing and over-exploitation of the reef resources in the East Asia-Pacific region were compounded by mass coral bleaching and consequent coral mortality. Coral bleaching events are increasing in frequency, extent and severity. These large-scale bleaching events are linked to global climate change and are expected to be a recurring problem. Because most island and coastal populations in the East Asia-Pacific region depend on coral reefs for nutrition, fisheries and tourist income, as well as coastal protection, mass coral bleaching and mortality create an environmental and socioeconomic crisis that requires development of a focused, coordinated response. This response must be based not only on an understanding of the causes of coral bleaching and the interaction of bleaching with other coral reef stressors, but also in recognition of initiatives already ongoing which seek to understand and mitigate the long-term impacts on coral reefs. In November 1998, the International Coral Reef Initiative issued a “Statement on Coral Bleaching,” anticipating negative consequences to tourism, fishing, coastal protection and other coral reef services. The Statement calls for interdisciplinary research to evaluate the immediate and ultimate causes of coral bleaching, and the effect of coral bleaching on the ecosystem. In 1999, a commitment was made by U.S. Agency for International Development and the U.S. Department of State, through the East Asia and Pacific Environmental Initiative (EAPEI), to research and address the socioeconomic impacts resulting from coral bleaching and to bring this information to

appropriate international forums. These goals were realized in part through a special session at the 9th International Coral Reef Symposium in Bali, Indonesia, supported by EAPEI funding. This compendium captures the symposium’s special session results. It brings together the experience in the science and management of coral reefs under conditions of climate change, emphasizing socioeconomic aspects of coral bleaching. We sincerely hope that this compendium will be both informative and interesting to practitioners, decisionmakers and researchers. We hope it focuses attention on the need for effective and innovative approaches to management of coral reefs. With awareness, knowledge and action, the viability of coral reefs will be assured. Seebauer, J. (2001) Zoology of Porites cylindrica: Potential for use in reef-rehabilitation transplantation efforts. Geneseo Journal of Science and Mathematics 2, 26-34. The life history strategies of a branching, arborescent coral, Porites cylindrica, were observed at Orpheus Island on the Great Barrier Reef, Australia, to investigate its potential for use for transplantation programs in reef rehabilitation efforts. My research suggests that Porites cylindrica is a tolerant, long-lived, fast-growing coral which relies heavily on asexual fragmentation as a mode of reproduction and dispersal. P. cylindrica demonstrated weak competitive strategies but was able to avoid most confrontations by using the fugitative strategy (Jackson 1979) to acquire living space in a direction away from a potential competitor. The data also suggest that P. cylindrica has taken deleterious incidents like breakage and turned them into advantageous events by reproducing asexually via fragmentation, possibly increasing its fitness. No differences in percent mortality were found due to colony size. The qualities of fast growth, ability to out-compete other coral species for living space, tolerance to environmental extremes, and resistance to bleaching allow P. cylindrica to be a major reef-building coral as well as a prime species for transplantation efforts. Shanks, AL, Grantham, BA and Carr, MH. (2003) Propagule dispersal distance and the size and spacing of marine reserves. Ecological Applications 13, S159-S169. This study compiled available information on the dispersal distance of the propagules of benthic marine organisms and used this information in the development of criteria for the design of marine reserves. Many benthic marine organisms release propagules that spend time in the water column before settlement. During this period, ocean currents transport or disperse the propagules. When considering the size of a marine reserve and the spacing between reserves, one must consider the distance which propagules disperse. We could find estimates of dispersal distance for 32 taxa; for 25 of these, we were also able to find data on the time the propagules spent dispersing. Dispersal distance ranged from meters to thousands of kilometers, and time in the plankton ranged from minutes to months. A significant positive correlation was found between the log-transformed duration in the plankton and the log-transformed dispersal distance (r=0.7776, r2=0.60, df=1.25. P=0.000); the more time propagules spend in the water column the further they tend to be dispersed. The frequency distribution of the log-transformed dispersal distance is bimodal (kurtosis=21.29, t=-4.062, P<0.001) with a gap between 1 and 20 km. Propagules

that dispersed ,1 km spent less time in the plankton (<100 h), or if they remained in the plankton for a longer period, they tended to remain in the waters near the bottom. Propagules that dispersed >20 km spent more than 300 h in the plankton. The bimodal nature of the distribution suggests that evolutionary constraints may reduce the likelihood of evolving mid-range dispersal strategies (i.e., dispersal between 1 and 20 km) resulting in two evolutionarily stable dispersal strategies: dispersal <1 km or .>~20 km. We suggest that reserves be designed large enough to contain the short-distance dispersing propagules and be spaced far enough apart that long-distance dispersing propagules released from one reserve can settle in adjacent reserves. A reserve 4–6 km in diameter should be large enough to contain the larvae of short-distance dispersers, and reserves spaced 10–20 km apart should be close enough to capture propagules released from adjacent reserves. Shenkar, N, Fine, M, Kramarsky-Winter, E and Loya, Y. (2006) Population dynamics of zooxanthellae during a bacterial bleaching event. Coral Reefs 25, 223–227. Each summer 80–90% of the colonies of Oculina patagonica undergo bleaching off the Mediterranean coast of Israel. To investigate fluctuations through a yearly bleaching cycle, monthly measurements of zooxanthella density, mitotic index and chlorophyll-a concentration were conducted. Results showed (1) a significant negative correlation between sea surface temperature (SST) and zooxanthella density; (2) both significantly lower zooxanthella mitotic index and higher chlorophyll-a per zooxanthella content during the bleaching season compared with the non-bleaching period; (3) prior to bleaching, a lag between the peak of zooxanthella density and chlorophyll-a concentration followed by a similar lag during recovery. Zooxanthella density declined significantly between March and May while chlorophyll-a concentration peaked in April, and then declined. Zooxanthella density increased significantly in November while chlorophyll-a concentration increased significantly in January. We conclude that during bacterial bleaching events, zooxanthellae are severely damaged. However, by the time of the following bleaching event the coral tissues regain their ‘‘normal’’ (pre-bleaching) zooxanthella population density. Sheppard, C. (1999) Coral mortality in the Chagos Archipelago. University of Warwick: Coventry (UK), 22 pp. Most of the corals and soft coral on seaward reefs of all six Chagos atolls have recently been killed. Reef condition is explained and illustrated with numerous photographs, and comparisons are made between the present situation (in 1999) and that found by earlier surveys, particularly during 1996. Today on seaward reefs, only about 12% of the substrate is live coral, 40% is dead coral identifiable as such, and 40% is unidentifiable dead substrate, much of which is almost certainly severely eroded, dead coral. In 1996, an average of about 75% of the substrate was living coral and soft corals. Some reefs have fared better than others, but on some, such as the seaward reef of Nelson Island, no living coral or soft coral (or any living coelenterate at all) was seen during the course of a 20 minute snorkel, in clear water, over these once spectacular reefs. Most lagoons contained greater amounts of living coral (about 28% living coral with 67% of the substrate being dead colonies or bare). The mortality here, therefore, is only about three quarters. Most corals are becoming heavily eroded, leaving

extensive rubble. Rubble on reefs is highly erosive and tends to prevent recolonisation. Reef fish were not systematically counted, but clearly on many reefs they too were less than 25% of their former abundance and diversity. The same applies to other invertebrate groups such as the starfish, urchins and molluscs which appear to have become very uncommon species in Chagos at least in the shallowest 20 m. This represents an unprecedented destruction and change to the coral reef community. Possible consequences are outlined and recommendations made. It is clear that this massive change requires a substantial re-think of present approaches and research priorities in coral reef conservation and science. Sheppard, C, Spalding, M, Bradshaw, C and Wilson, S. (2002) Erosion vs recovery of coral reefs after 1998 El Niño: Chagos reefs, Indian Ocean. Ambio 31, 40-48. Three years after most corals died on the central Indian Ocean reefs of Chagos, erosion and recovery were studied to 30 m depth. Mortality was near-total to 15 m deep in northern atolls, and to > 35 m in central and southern atolls. Some reef surfaces have ‘dropped’ 1.5 m due to the loss of dense coral thickets. Coral bioerosion is substantial, reducing 3-D reef ‘structure’ and forming unconsolidated rubble. Juvenile corals are abundant, though mostly on eroding or unstable substrates, and are of less robust species. Reef fish abundance and diversity at 15 m depth remains high; species dependent on corals have diminished, while some herbivores and detritivores have increased. A new sea surface temperature (SST) data set shows that mean SST has risen 0.65°C since 1950. The critical SST causing the mortality in Chagos was 29.9°C. Sheppard, C. (2003) Predicted recurrences of mass coral mortality in the Indian Ocean. Nature 425, 294-297. In 1998, more than 90% of shallow corals were killed on most Indian Ocean reefs. High sea surface temperature (SST) was a primary cause, acting directly or by interacting with other factors. Mean SSTs have been forecast to rise above the 1998 values in a few decades; however, forecast SSTs rarely flow seamlessly from historical data, or may show erroneous seasonal oscillations, precluding an accurate prediction of when lethal SSTs will recur. Differential acclimation by corals in different places complicates this further. Here I scale forecast SSTs at 33 Indian Ocean sites where most shallow corals died in 1998 (ref. 1) to identify geographical patterns in the timing of probable repeat occurrences. Reefs located 10–15° south will be affected every 5 years by 2010–2025. North and south from this, dates recede in a pattern not directly related to present SSTs; paradoxically, some of the warmest sites may be affected last. Temperatures lethal to corals vary in this region by 6°C, and acclimation of a modest 2°C by corals could prolong their survival by nearly 100 years.

Sheppard, C and Obura, D. (2004) Corals and reefs of Cosmoledo and Aldabra atolls: Extent of damage, assemblage shifts and recovery following the severe mortality of 1998. Journal of Natural History 39, 103-121. An updated list of over 200 species of corals from Cosmoledo and Aldabra atolls is presented, which more than doubles previously known species diversity, and establishes these atolls as amongst the most species-rich in the Western Indian Ocean. However, partly this is an artefact of a new method of recording with digital cameras, described here, which greatly improves recording efficiency. This is the first underwater study of Cosmoledo, and the first for Aldabra outside the expedition reports cited. The survey extended to .30 m depth, and comes after the 1998 massive coral mortality. Coral cover was virtually eliminated at that time to about 8-10 m depth in Cosmoledo on seaward slopes, below which coral mortality was only about 50%. Mortality was selective regarding different species, genera and families. Cosmoledo’s lagoon of .150 km2 is shallower than the ‘critical depth’ of 8–10 m, resulting in an almost complete elimination of corals. To compare these atolls with other reefs in the region, critical depths are summarized for over 25 Indian Ocean locations. New coral recruits are abundant in the shallows of Cosmoledo and Aldabra 4 years later, though cover remains very low. Much bare rock remains (with turf algae) and some genera such as Acropora, previously apparently abundant, remain relatively very scarce. Apart from Porites, whose higher survival is now well documented, the best survivors from the 1998 mortality, and the most successful recruitment of new corals, are of faviids. Soft corals remain extremely scarce in all locations examined above the ‘critical depth’. It is predicted that there may be a shift in the identity of the main species of corals in these atolls for many years. Siebeck, UE, Marshall, NJ, Klüter, A and Hoegh-Guldberg, O. (2006) Monitoring coral bleaching using a colour reference card. Coral Reefs 25, 453-460. Assessment of the extent of coral bleaching has become an important part of studies that aim to understand the condition of coral reefs. In this study a reference card that uses differences in coral colour was developed as an inexpensive, rapid and non-invasive method for the assessment of bleaching. The card uses a 6 point brightness/saturation scale within four colour hues to record changes in bleaching state. Changes on the scale of 2 units or more reflect a change in symbiont density and chlorophyll a content, and therefore the bleaching state of the coral. When used by non-specialist observers in the field (here on an intertidal reef flat), there was an inter-observer error of ± 1 colour score. This technique improves on existing subjective assessment of bleaching state by visual observation and offers the potential for rapid, wide-area assessment of changing coral condition. Smith, L and Birkeland, C. (2003) Managing NPSA’s coral reefs in the face of global warming. University of Hawaii: Honolulu (USA), 32 pp. Shallow pools in Ofu Lagoon in the National Park of American Samoa vary in thermal conditions, with temperatures in some pools reaching 35 °C and fluctuating daily by >6 °C. Yet the pools support diverse coral communities, including many species thought to be sensitive to high temperatures such as Acropora, Pocillopora, and Millepora spp. A one year experiment was carried out to evaluate the survival and growth of corals transplanted from cooler to warmer pools and vice versa. Nubbins of

Pocillopora damicornis and Porites cylindrica were obtained from three sites in Ofu Lagoon and one site in Ofu Harbor, dyed with Alizarin red S for measuring growth, and transplanted between the sites. The Ofu Lagoon sites varied in daily maximum temperatures and daily temperature ranges, and Ofu Harbor had more frequent warm temperatures and greater daily ranges than the Ofu Lagoon sites. Water temperatures were monitored hourly at the four sites during the one year period, and other water quality and physical characteristics were studied at the lagoon sites. Regardless of site of origin, both species had the greatest growth in Pool 300, the lagoon pool with the highest daily maximum temperatures and greatest daily temperature ranges of the three lagoon sites. Pool 300 is also the shallowest of the three lagoon sites, thus likely subject to the greatest irradiance. These results do not support the hypothesis that corals found in Ofu Lagoon pools with relatively high temperatures are better acclimatized or adapted to high temperatures than corals in the cooler portions of the lagoon. The hypothesis that corals found in Ofu Lagoon are better acclimatized or adapted to high temperatures than corals outside the lagoon in deeper water has not yet been tested. Extrinsic factors such as water motion, dissolved oxygen, or habitat characteristics may explain the greater survival and growth of coral transplants in Pool 300 than in the other lagoon pools. Water motion is known to increase the ability of corals to withstand high temperatures and irradiance, and flow velocity was higher in Pool 300 than the other two lagoon pools. Hyperoxia, or dissolved oxygen saturation >150%, stimulates the metabolism of zooxanthellae, and hyperoxia was most frequent and highest in Pool 300. Relatively low turbidity during stormy weather was found in Pool 300, which also had the least sandy substrate, possibly limiting sedimentation and abrasion of corals. Extrinsic factors may be an important consideration in establishing Marine Protected Areas to mitigate the effects of global warming on coral communities. Smith, L. (2004) Influence of water motion on resistance of corals to high temperatures: Evidence from a field transplant experiment. University of Hawaii: Honolulu (USA), 19 pp. Pocillopora damicornis and Porites cylindrica were transplanted between sites within the backreef moat on Ofu Island, American Samoa, to investigate the effects of extrinsic factors such as water motion on resistance of corals to high temperatures. Coral transplants were obtained from a relatively deep Source Pool, dyed with Alizarin red S to measure linear extension, and transplanted into two shallower pools, Test Pool 1 and Test Pool 2, as well as back into the Source Pool. Test Pool 1 had the highest daily maximum temperatures, the greatest duration of high temperatures, and the greatest daily fluctuations of temperatures of the three study sites. However, both transplanted species grew more in Test Pool 1 than the other two sites. Mean flow speeds were highest in Test Pool 1, especially on adjacent reef flats. Transplants in Test Pool 1 were located only 1-5 m from the reef flat, whereas in the other two pools they were located >35 m from the reef flat, thus transplants in Test Pool 1 were more exposed to high flow speeds from the reef flat than the transplants in the other two pools. The greater coral transplant growth rates in Test Pool 1 suggest that water motion enhanced the resistance of the transplants to the relatively high temperatures at this site.

Smith, DJ, Suggett, DJ and Baker, NR. (2005) Is photoinhibition of zooxanthellae photosynthesis the primary cause of thermal bleaching in corals? Global Change Biology 11, 1–11. The bleaching of corals in response to increases in temperature has resulted in significant coral reef degradation in many tropical marine ecosystems. This bleaching has frequently been attributed to photoinhibition of photosynthetic electron transport and the consequent photodamage to photosystem II (PSII) and the production of damaging reactive oxygen species (ROS) in the zooxanthellae (Symbiodinium spp.). However, these events may be because of perturbations of other processes occurring within the zooxanthellae or the host cells, and consequently constitute only secondary responses to temperature increase. The processes involved with the onset of photoinhibition of electron transport, photodamage to PSII and pigment bleaching in coral zooxanthellae are reviewed. Consideration is given to how increases in temperature might lead to perturbations of metabolic processes in the zooxanthellae and/or their host cells, which could trigger events leading to bleaching. It is concluded that production of ROS by the thylakoid photosynthetic apparatus in the zooxanthellae plays a major role in the onset of bleaching resulting from photoinhibition of photosynthesis, although it is not clear which particular ROS are involved. It is suggested that hydrogen peroxide generated in the zooxanthellae may have a signalling role in triggering the mechanisms that result in expulsion of zooxanthellae from corals. Smith, JE, Shaw, M, Edwards, RA, Obura, D, Pantos, O, Sala, E, Sandin, SA, Smriga, S, Hatay, M and Rohwer, FL. (2006) Indirect effects of algae on coral: algae-mediated, microbe-induced coral mortality. Ecology Letters 9, 835-845. Declines in coral cover are generally associated with increases in the abundance of fleshy algae. In many cases, it remains unclear whether algae are responsible, directly or indirectly, for coral death or whether they simply settle on dead coral surfaces. Here, we show that algae can indirectly cause coral mortality by enhancing microbial activity via the release of dissolved compounds. When coral and algae were placed in chambers together but separated by a 0.02 mum filter, corals suffered 100% mortality. With the addition of the broad-spectrum antibiotic ampicillin, mortality was completely prevented. Physiological measurements showed complementary patterns of increasing coral stress with proximity to algae. Our results suggest that as human impacts increase and algae become more abundant on reefs a positive feedback loop may be created whereby compounds released by algae enhance microbial activity on live coral surfaces causing mortality of corals and further algal growth. Smith-Keune, C and van Oppen, M. (2006) Genetic structure of a reef-building coral from thermally distinct environments on the Great Barrier Reef. Coral Reefs 25, 493–502. Adaptation to localised thermal regimes is facilitated by restricted gene flow, ultimately leading to genetic divergence among populations and differences in their physiological tolerances. Allozyme analysis of six polymorphic loci was used to assess genetic differentiation between nine populations of the reef-building coral Acropora millepora over a latitudinal temperature gradient on the inshore regions of

the Great Barrier Reef (GBR). Small but significant genetic differentiation indicative of moderate levels of gene flow (pairwise FST 0.023 to 0.077) was found between southern populations of A. millepora in cooler regions of the GBR and the warmer, central or northern GBR populations. Patterns of genetic differentiation at these putatively neutral allozyme loci broadly matched experimental variation in thermal tolerance and were consistent with local thermal regimes (warmest monthly-averages) for the A. millepora populations examined. It is therefore hypothesized that natural selection has influenced the thermal tolerance of the A. millepora populations examined and greater genetic divergence is likely to be revealed by examination of genetic markers under the direct effects of natural selection. Soong, K and Chen, T-A. (2003) Coral transplantation: Regeneration and growth of Acropora fragments in a nursery. Restoration Ecology 11, 62-71. Coral reef degradation has been widely reported for the past 20 years. Because the recovery rate is usually low, various methods of restoration have been explored in different regions of the world. Among the effective and commonly used methods to restore coral communities is the transplantation of coral colonies or fragments. In this investigation fragments of Acropora pulchra were used in a semiprotected nursery in southern Taiwan between 1996 and 1998 to test, in situ, the possible effects of different factors on the generation of new branches and the initial skeletal extension rates of transplants. The variables under study here were the origin and length of the fragments, their new orientation, presence of tissue injury, and position in the fragment. All these factors were found to make a difference in either one or both aspects of coral growth (i.e., branching frequency and skeletal extension rate). These two factors clearly determine the success rate of a small fragment developing into a large colony that has a much higher probability to survive and grow on its own. It is now obvious that the efficiency of coral generation through fragment culture can be enhanced if the variables examined here are taken into consideration. Once coral colonies are formed, they can be fragmented again to generate more corals or can be transplanted to a suitable site. Soong, K, Chen, M-H, Chen, C-L, Dai, D-F, Fan, T-Y, Li, J-J and Fan, H. (2003) Spatial and temporal variation of coral recruitment in Taiwan. Coral Reefs 22, 224-228. The effect of recruitment on the structure and dynamics of marine populations and communities has received much attention in recent years (e.g., Caley et al. 1996). Even for long-lived corals, recruitment plays a critical role in the distribution and abundance of species (Connell et al. 1997). Most research on coral recruitment has been conducted in the last three decades (e.g., Birkeland 1977; Hughes et al. 1999). Research has now shown that coral larvae are sensitive to microhabitats, e.g., substrate composition (Benayahu and Loya 1984), neighbors (Maida et al. 1995), light conditions, and orientation of substrates (Babcock and Mundy 1996). Densities and composition of coral recruits also differ among seasons and between years, often by several orders of magnitude (e.g., Wallace 1985). The patterns of recruitment are known to change with different spatial scales (Dunstan and Johnson 1998). The scale of spatial distribution of coral recruitment has been explored mostly on the Great Barrier Reef (GBR) and its neighboring reefs (e.g., Hughes et al. 1999). It has been suggested that the pattern found in this 2,000-km stretch differs from

reefs that are relatively isolated from one another (Harriott and Banks 1995). For example, a larva in the GBR has a good chance, after drifting for 2 weeks and for 200 km, to settle on the GBR. This scale of self-seeding is unlikely for many other reefs that are relatively isolated and small (Kojis and Quinn 2001). In this study, we report on recruitment rates in Taiwan and, in particular, spatial variation in recruitment within and between areas in Taiwan. Sotka, EE and Thacker, RW. (2005) Do some corals like it hot? Trends in Ecology and Evolution 20, 59-62. Global increases in sea temperatures threaten coral reef resilience because thermal stress can cause corals to bleach; that is, to lose their photosynthetic microalgal symbionts. Recent evidence suggests that some corals associate with genotypes of microalgae that resist future thermal stress, however, these genotypes might provide less energy for growth when thermal stresses are curtailed. Coral reef resilience depends on whether phenotypic and genotypic changes in host–symbiont associations can match projected increases in the frequency and severity of thermal stress, as well as on our ability to ameliorate continuing human impacts. Soto, CG. (2002) The potential impacts of global climate change on Marine Protected Areas. Reviews in Fish Biology and Fisheries 11, 181-195. The potential effects of global climate change on marine protected areas do not appear to have been addressed in the literature. This paper examines the literature on protected areas, conservation biology, marine ecology, oceanography, and climate change, and reviews some of the relevant differences between marine and terrestrial environments. Frameworks and classifications systems used in protected area design are discussed. Finally, a framework that summarizes some of the important oceanographic processes and their links to the food chain are reviewed. Species abundance and distribution are expected to change as a result of global climate change, potentially compromising the efficacy of marine protected areas as biodiversity conservation tools. This review suggests the need for: further interdisciplinary research and the use of linked models; an increase in marine protected areas for biodiversity conservation and as research sites for teasing apart fishing effects from climate effects; a temporally responsive approach to siting new marine protected areas, shifting their locations if necessary; and large-scale ecosystem/integrated management approaches to address the competing uses of the oceans and boundary-less threats such as global climate change and pollution. Souter, DW and Lindén, O. (2000) The health and future of coral reef systems. Ocean and Coastal Management 43, 657-688. Coral reefs are among the most productive and diverse ecosystems on earth and provide a multitude of valuable ecosystem services. Moreover, the resources derived from coral reefs are essential to the food security of millions of people living within tropical coastal communities. Unfortunately, burgeoning human populations in coastal regions are placing an unsustainable burden on these resources such that degradation of coral reefs is common. In addition, during 1998, El Niño driven increases in sea temperature caused a mass bleaching event that further degraded many of the world's coral reefs. This article provides a brief review of the status of the

world's coral reefs and highlights their value to society. Also, the anthropogenic and natural disturbances that threaten the future of coral reefs are discussed and finally, this article offers some potential remedies that promote sustainable use of coral reef resources thus ensuring their future survival. Souter, DW, Obura, DO and Lindén, O (eds.) (2000) Coral reef degradation in the Indian Ocean: Status report 2000. CORDIO: Stockholm (Sweden), 205 pp. The World Conservation Strategy (IUCN/UNEP/WWF) identifies coral reefs as one of the essential global life support systems necessary for food production, health and other aspects of human survival and sustainable development. However, their future is being threatened by overexploitation, coastal development, landbased pollution and moreover, by global climate change. The last two decades has been the warmest period ever recorded during which, 1998 was the hottest single year. As a consequence, the mean temperatures of the world’s oceans are increasing at an accelerating rate which, at present, is approximately 2°C per century. In addition, 1998 witnessed the strongest El Niño ever recorded. The long period for which this El Niño prevailed and its extreme magnitude exacerbated already increased sea-surface temperatures. In 1998, as a consequence of these El Niño driven increases in sea temperature, corals throughout the world’s tropical oceans suffered the most severe mass bleaching event ever reported. As a direct response to the massive degradation caused by this bleaching event, the CORDIO Program (Coral Degradation in the Indian Ocean) was initiated early in 1999. CORDIO’s objectives are to determine the long-term impacts of degradation of coral reefs on coastal communities and national economies. Research under CORDIO focuses on: • the bio-physical impacts of coral degradation as a result of bleaching and other disturbances, and the long term prospects of recovery, • the socio-economic impacts of coral mortality and options for mitigating these through management and development of alternative livelihoods, • the prospects of restoration and rehabilitation of reefs to accelerate their ecological and economic recovery This second annual status report follows on from the first report that was based on initial assessments of coral mortality and presents the results of CORDIO’s first full year of monitoring and research. BIOPHYSICAL EFFECTS Monitoring of the biophysical effects of the El Niño was conducted in 11 countries in East Africa (Kenya, Tanzania, Mozambique), South Asia (Sri Lanka, Maldives, India), and the central Indian Ocean islands (Seychelles, Mauritius, Reunion, Comoros, Madagascar). Reef degradation was most pronounced on the reefs of mainland East Africa and on the islands of Socotra, Comoros, Mayotte, Seychelles, and the islands of South Asia. In these areas coral mortality generally exceeded 50% and reached 95% or more in many locations. Large areas of coral reefs in the central and western Indian Ocean have undergone a total transformation where the former coral reef is now covered by thick carpets of filamentous algae. Areas that suffered minimal bleaching and mortality included the more southerly islands of Mauritius and Reunion, where cyclonic activity retarded increases in sea-surface temperature. Also, in other areas, such as western Unguja and Pemba Islands and on some reefs of the Seychelles Bank, upwelling of cooler waters from off the continental shelf and bank slopes moderated increased sea surface temperatures and prevented the subsequent bleaching of corals.

No sites were completely unaffected by bleaching. Although bleaching and mortality among corals was observed at depths between 40 m and 50 m, generally the damage was much greater in shallow waters. Corals growing in shallow areas with restricted water circulation are generally considered to be more tolerant of high sea temperatures than corals situated in deeper waters. However, the magnitude and duration of increases in sea temperature caused by the 1998 El Niño event overwhelmed any capacity possessed by corals growing in shallow water to tolerate such conditions. Subsequently, it was these corals that suffered disproportionately high mortality. Soon after, coral competitors (algae and some zoanthids (“false corals”)) began to occupy space made available by the massive coral mortality. Initial investigations of the fish communities associated with the reefs indicated that herbivorous species increased in abundance while the abundance of corallivorous species declined. In several areas (Maldives, Seychelles, Chagos, Socotra) the dead coral skeletons have been broken down into rubble and no longer prevent oceanic waves from penetrating into sheltered waters in the lee of the reef. Subsequently, erosion of the beaches in these areas appears to have increased. RECOVERY Coral recovery is expected to occur through re-growth and/or fragmentation of colonies that survived the mortality event and also through recruitment of planula larvae from the water column. With the exception of some reefs of Maldives, recovery through the recruitment of larvae has been minimal. Re-growth of surviving corals has occurred on some reefs, but generally has not been significant. The degree to which recovery of reefs will be mediated by the influx of coral larvae from more distant reefs is currently unknown largely because so few reefs have escaped the mass bleaching event relatively unscathed and could potentially serve as sources of larvae. Priority areas for biophysical research for the year 2000 include: identification of source reefs (reefs from which a supply of coral larvae could originate) and identification of threats to these reefs, assessment of processes of recovery and degradation of reefs in affected areas, the affect that local species extinction exerts on the structure of recovering reef communities with respect to dispersal and influx of larvae, and the promotion of regional synoptic research projects. SOCIO-ECONOMIC EFFECTS Socio-economic monitoring was conducted in seven countries (Madagascar, Comoros, Tanzania, Kenya, Maldives, Sri Lanka, India). Reef fisheries have not yet shown any obvious declines attributable to the El Niño induced coral bleaching. This is probably attributable to the fact that, at present, the structural integrity of many of these reefs remains largely intact and continues to provide the structure needed by fish communities. However, as soon as the reef framework becomes degraded by erosion, a decline in fish catches is expected. Because a substantial proportion of the coastal fishery is subsistence fishing, the impact of declining fish stocks may threaten food security of a large proportion of the coastal population in each of these countries. Tourist divers interviewed at Zanzibar, Mombasa and Maldives indicated the state of the reef would affect their choice of destination and whether to dive, resulting in significant estimated economic and financial losses. However, differences in the tourism market and product among destinations will greatly affect their ability to offer alternative activities to preserve income. The principal questions derived from these investigations identify the importance of links between the biophysical and socio-

economic environments and the need to adapt both the fisheries and tourism industries accordingly. PROGRAM FOR YEAR 2000 CORDIO held its second annual workshop, “Coral Reef Degradation in the Indian Ocean (CORDIO): Progress to Date and Directions for 2000 and Beyond” between February 10th and 12th, 2000, in Lamu, Kenya. Thirtyone participants attended the workshop to review status of coral reefs in the region, and the progress of research to date. Working group discussions were held in five areas: 1. Climate change and long term trends 2. Regional research 3. Socio-economic considerations 4. Management and awareness issues 5. Database requirements The conclusions of the working groups are included in this report. Of particular importance were discussions held on the novel challenges to research and management posed by climate change and the false vision of a pristine historical reef state that has underpinned most current research. What is needed is almost an emerging science that develops innovative ways to measure and interpret the influence of climate change on ecological systems. Of utmost importance is the compilation of long-term records (e.g. meteorological data) to establish long-term trends, measurement of simple variables (such as beach width) over large spatial scales to establish synoptic baselines and greater dedication to monitoring key processes such as recruitment rather than gross variables such as coral cover. The increasing frequency and severity of mass coral bleaching events is the single most dramatic and tangible demonstration of global climate change. Predictions of the consequences of gradual rises in temperature and sea level have been eclipsed by the effects of violent fluctuations of natural climatic cycles. El Niño events of a magnitude previously experienced only once every 100 years are now occurring at intervals ranging between three and 20 years, with severe impacts extending beyond their original limits within the Eastern Pacific to the equatorial and monsoon regions of the Indian Ocean. The development of alternative livelihoods for subsistence fishermen and others dependent on reef resources is an important area for research and development. Aquaculture of commercially valuable organisms, expansion of marine resource use to include a wider variety of species thus alleviating the pressure on target species and the development of terrestrial opportunities to improve livelihoods in terms of income and/or food security is desirable. However, participation of all stakeholders in the process of development and implementation is paramount for the success of such schemes Coral reefs are among the first ecosystems on the planet that are being dramatically affected by global climate change. Their high biodiversity and their fundamental importance to coastal populations raises the need for problem solving research and management. Bleaching of coral as a result of increased sea temperatures caused by global climate change offers a unique opportunity to investigate the effects of climate change on an ecosystem scale and within a relatively small time frame. Conseqvently, CORDIO is establishing a process for addressing this global challenge in the western Indian Ocean. Despite the growing weight of local and global threats, CORDIO, in 2000 and 2001, will attempt to improve the fate of coral reefs through research and development projects that support management and improve the livelihoods of people dependent on coral reef ecosystems.

Souter, DW and Lindén, O (eds.) (2005) Coral reef degradation in the Indian Ocean: Status report 2005. Cordio: Kalmar (Sweden), 285 pp. The assessments of 50 contributing authors focusing on coral reefs and related coastal ecosystems, and dependent communities in 9 countries in South Asia and the central and western Indian Ocean report that: Status of Coral Reefs ● The bio-physical condition of many reefs continues to decline. ● Recovery from bleaching associated coral mortality is generally slow and patchy with widespread changes in species composition of adult and juvenile coral communities. ● Recovery is more rapid on reefs that are situated within managed areas or are remote from the influence of other human disturbances. ● Recovery in areas subject to human influences has been retarded. ● The primary causes of coral reef degradation are: � Bleaching, which is occuring more frequently and has accelerated the degradation caused by: � Overexploitation of fish and of other organisms on reefs throughout the region; � Destructive fishing, which become an increasing problem as fish stocks decline; due to lack of enforcement destructive fishing has destroyed reefs in formally protected areas (MPA’s) during the last couple of years; � Pollution and sedimentation, mainly from landbased human activities. ● The impacts of the tsunami of 26th December 2004 on coral reefs was highly variable and ranged from negligible (Gulf of Mannar, India, Maldives, East Africa) to moderate (parts of Andaman and Nicobar Islands) to extreme (parts of Sri Lanka, Nicobar Islands). ● The primary factors determining the severity of damage caused by the tsunami were: � How exposed a reef was to the direct force of the wave; � The local bathymetry surrounding a reef; � The geological composition of a reef; � The condition of the reef; reefs that had suffered extensive coral mortality as a result of the 1998 El Niño were more vulnerable to the force of the tsunami. ● The reasons for continued coral reef degradation are: � High dependence on coral reef resources as a result of few alternative sources of food and income; � Open access; fishing and exploitation of other coastal resources are unregulated in many countries; as a result, unemployment and a lack of opportunities elsewhere in society are directly linked with coastal degradation; � Low awareness of the importance of healthy coastal ecosystems and the impacts of human activities; � Inadequate laws and regulations; � Poor enforcement of existing laws and regulations; � Responsabilities dispersed among several agencies and a lack of coordination between these agencies; � Insufficient political will to strengthen laws and regulations and improve enforcement. Actions Necessary to Achieve Sustainable Use of Coral Reef Resources Research and Monitoring

● Continue to strengthen environmental monitoring programmes in each country so that management decisions are made using the best available information and so that management strategies can be adapted to cope with changing situations. ● Engage communities in monitoring and management in order to raise awareness of the importance of coral reefs, establish behaviours that are not environmentally degrading and secure community support for sustainable resource use. ● Continue to develop research capacity in the region to specifically address management issues. Management and Governance ● Increase the area of coral reefs currently under management and incorporate management strategies within the broader frameworks of integrated coastal zone and catchment management in order to effectively deal with land-based issues such as pollution, sedimentation and coastal development. ● Ensure participation of all stakeholders in the development and revision of management strategies in order to open channels of communication and achieve ownership and transparency of process. ● Improve management capacity among local and national authorities. Years of inactivity, often deliberate, must be stopped. In many cases the roots of the problem is at the political level where deeprooted cronyism, unwillingness to follow laws and regulations because key voter groups may be affected, and outright corruption prevents responsible behaviour. ● Strengthen laws and regulations where necessary. Clarify the responsabilities of different agencies. ● Enforcement of laws and regulations must be strengthened. Punishments against destructive behaviour must be such that they are genuine disincentives. ● Improve enforcement by providing greater manpower, equipment and financial resources. ● Encourage community-based protection and enforcement in partnership with government agencies by devolving to stakeholders the responsibility for direct conservation of resources and enforcement of laws. ● Strengthen political will through the education of decision makers of the tangible benefits of management. AlternativeLivelihoods and Income Diversification ● Stimulate the introduction of income diversification and alternative livelihood schemes that are environmentally sustainable and economically viable and socially acceptable. Examples of such activities can be found in areas related to agriculture, aquaculture, waste management, tourism and manufacturing. A wide range of activities should be encouraged in order to ensure long-term viability. ● If necessary, establish financial mechanisms targeted specifically at improving the economic development of the poor. ● If necessary, secure government support for alternative livelihood and income diversification schemes. Education and Awareness ● Raise awareness of the importance of healthy coastal ecosystems and the impacts of various human activities through: � the introduction of marine studies in school curricula; � community-based monitoring and management activities; � awareness raising programmes targeting specific sectors of society; � the use of mass media.

Spalding, MD and Jarvis, GE. (2002) The impact of the 1998 coral mortality on reef fish communities in the Seychelles. Marine Pollution Bulletin 44, 309-321. Coral reef fish communities in the Seychelles are highly diverse and remain less affected by the direct impacts of human activities than those on many other coral reefs in the Indian Ocean. These factors make them highly suitable for a detailed survey of the impacts of the 1998 mass coral mortality, which devastated the coral faunas of the region. Using underwater visual census (UVC) techniques, fish communities were sampled in three localities in the southern Seychelles and one locality in the northern (granitic) Seychelles. Initial surveys were undertaken from the latter site in 1997. Surveys were undertaken at all sites during the coral bleaching episode in 1998 prior to any major changes in the reef fish communities. Repeat surveys were undertaken in 1999 one year after the coral mortality. Over 250 fish species were sampled from 35 families. Results suggest that changes in the overall fish community structures are not great, despite massive changes in the benthic cover. Significant changes have been observed in a number of individual species. These include those most heavily dependent on live coral cover for shelter or sustenance. Future potential changes are discussed, and potential management interventions are considered. Spencer, T, Teleki, KA, Bradshaw, C and Spalding, MD. (2000) Coral bleaching in the southern Seychelles during the 1997-1998 Indian Ocean warm event. Marine Pollution Bulletin 40, 569-586. Coral bleaching shows complex spatial-temporal dynamics at several scales. Recent developments in ocean surface remote sensing technologies allow for both a better appreciation of these dynamics and an opportunity to place more local studies of coral bleaching and bleaching-related coral mortality into wider oceanographic and biogeographic contexts. Coral bleaching is described at four coral reefs in the southern Seychelles (sea area 6-10°S 45-54°E) during March-May 1998. Bleaching intensity varied between locations, between environments at the within-reef scale, and between coral growth forms. These data are compared with bleaching reports and sea surface temperature statistics for eight further stations in the western Indian Ocean to establish a link between bleaching and the unprecedented warming of the Indian Ocean during 1997/98. Implications for long-term reef history, and coral reef futures, in the western Indian Ocean are discussed. Sprecher, SG, Galle, S and Reichert, H. (2003) Substrate specificity and juvenile Faviid predominance of coral colonization at the Maldive Islands following the 1998 bleaching event. Coral Reefs 22, 130-132. During the coral bleaching event of 1998 in the Indopacific, there was extensive coral mortality in some areas (Naeem et al. 1998; Edwards et al. 2001; McClanahan et al. 2001; Mumby et al. 2001). For example, observations on North Male Atoll, the Maldives Islands, indicated that up to 90% of hermatypic corals died following the bleaching event. Prior to the event, 42% of the reef surface was covered with living hermatypic corals, while only 2% was covered by living corals after the event (Naeem et al. 1998; Edwards et al. 2001). Two substrates for coral colonization became available in great quantity as a result of coral mortality in 1998: first, numerous coral fragments that resulted from disintegration of dead Acroporidae and other branching

corals and, second, the surface of dead, massive corals. Here, we report on colonization of these two substrate types by juvenile corals 3 years after the bleaching event. Stewart, HL, Holbrook, SJ, Schmitt, RJ and Brooks, AJ. (2006) Symbiotic crabs maintain coral health by clearing sediments. Coral Reefs 25, 609-615. Stony corals are the foundation of coral reef ecosystems and form associations with other reef species. Many of these associations may be ecologically important and play a role in maintaining the health and diversity of reef systems, rendering it critical to understand the influence of symbiotic organisms in mediating responses to perturbation. This study demonstrates the importance of an association with trapeziid crabs in reducing adverse effects of sediments deposited on corals. In a field experiment, mortality rates of two species of branching corals were significantly lowered by the presence of crabs. All outplanted corals with crabs survived whereas 45–80% of corals without crabs died within a month. For surviving corals that lacked crabs, growth was slower and tissue bleaching and sediment load were higher. Laboratory experiments revealed that corals with crabs shed substantially more of the sediments deposited on coral surfaces, but also that crabs were most effective at removing grain sizes that were most damaging to coral tissues. The mechanism underlying this symbiotic relationship has not been recognized previously, and its role in maintaining coral health is likely to become even more critical as reefs worldwide experience increasing sedimentation. Stimson, J, Sakai, K and Sembali, H. (2002) Interspecific comparison of the symbiotic relationship in corals with high and low rates of bleaching-induced mortality. Coral Reefs 21, 409-421. Some coral species are more resistant than others to environmental factors that cause bleaching and bleaching-related mortality. This study compared aspects of the coral/zooxanthellae symbiosis in species and genera that suffered either high or low mortality during a bleaching event. These characteristics were assessed in Okinawa between March and June 1999, 5–10 months after the bleaching event there in August–September 1998. Species with low mortality rates generally had higher densities of zooxanthellae per square centimeter and a very low rate of release of degraded zooxanthellae. Low-mortality species also had more total coral tissue per square centimeter of coral surface area. The size of zooxanthellae varied little among species. The differences in these characteristics among coral species suggest that the symbiotic relationship operates very differently among coral species. Strychar, KB, Coates, M, Sammarco , PW and Piva, TJ. (2004) Bleaching as a pathogenic response in scleractinian corals, evidenced by high concentrations of apoptotic and necrotic zooxanthellae. Journal of Experimental Marine Biology and Ecology 304, 99-121. The study of symbiont cells lost from bleached scleractinian corals Acropora hyacinthus, Favites complanata, and Porites solida and octocorals Sarcophyton ehrenbergi, Sinularia sp., and Xenia sp. using flow cytometry shows that Symbiodinium die from either apoptosis or necrosis. Despite the majority of lost Symbiodinium cells being viable at 28°C, the predominance of apoptotic and necrotic

symbiont cells at higher temperatures indicates that the proportion of live cells decreases with increasing temperature. This implies that reinfection of corals at high temperatures by Symbiodinium lost from scleractinian corals may be less frequent than previously described, since many of the symbiont cells exhibit nonreversible symptoms of approaching cell death. The fraction of viable Symbiodinium cells lost from S. ehrenbergi, Xenia sp., and Sinularia at 32°C was greater than that at 28°C. At 34°C, the fraction of viable cells lost from S. ehrenbergi and Xenia sp. fell but not from Sinularia sp., which suggests that their symbionts have higher temperature tolerances. Thus, Symbiodinium from octocorals may represent ‘‘pools’’ of genetically resistant symbionts available for reinfection of other reef organisms. This has been proposed previously for Symbiodinium in some scleractinian corals, but this is the first evidence for such, particularly for an octocoral. Many of the viable cells, determined using Trypan blue staining techniques, are in fact actually undergoing apoptosis or necrosis, when examined using Annexin V-fluor and propidium iodide staining profiles. The characterization of more apoptotic and necrotic cells than viable cells is critical, as this indicates that the loss of Symbiodinium cells cannot be beneficial to other bleached corals for symbiotic reassociation. Suefuji, M and van Woesik, R. (2001) Coral recovery from the 1998 bleaching event is facilitated by Stegastes (Pisces: Pomacentridae) territories, Okinawa, Japan. Coral Reefs 20, 385-386. Abstract not available. Sydney Centre for International and Global Law. (2004) Global climate change and the Great Barrier Reef: Australia’s obligations under the World Heritage Convention. University of Sydney: Sydney (Australia), 39 pp.

1. We have been asked to provide a report in relation to Australia’s obligations under international law to protect the Great Barrier Reef World Heritage Area (‘GBRWHA’) from the effects of global climate change. In this connection we have been requested to determine whether Australia has complied with the 1972 Convention Concerning the Protection of the World Cultural and Natural Heritage

(‘World Heritage Convention’). 2. In relation to the World Heritage Convention we have been asked: • Whether Australia’s decision not to ratify the 1997 Kyoto Protocol

to the 1992 United Nations Framework Convention on Climate Change

amounts to a breach of Australia’s obligations under the World Heritage Convention in relation to the GBRWHA; • Whether Australia’s actions during the negotiations for the conclusion of the Kyoto Protocol amounted to a breach of Australia’s obligations under the World Heritage Convention in relation to the GBRWHA; • Whether a failure by Australia to commit to “deep cuts” in greenhouse gas emissions amounts to a breach of Australia’s obligations under the World Heritage Convention in relation to the GBRWHA; • Whether State Parties other than Australia have obligations pursuant to the World Heritage Convention to protect and conserve the GBRWHA and therefore whether the failure of another State to take action to reduce greenhouse gas emissions amounts to a breach of Article 6 of the World Heritage Convention;

• Whether Australia’s climate change policies come within the scope of the World Heritage Convention and therefore are required to be reported to the General Conference of the United Nations Educational, Scientific and Cultural Organization (‘UNESCO’) pursuant to Article 29(1) of the World Heritage Convention; and • Whether, and in what circumstances, the GBRWHA may be included on the List of World Heritage in Danger pursuant to Article 11 of the World Heritage Convention

Takahashi, S, Nakamura, T, Sakamizu, M, van Woesik, R and Yamasaki, H. (2004) Repair machinery of symbiotic photosynthesis as the primary target of heat stress for reef-building corals. Plant Cell Physiology 45, 251-255. In a coral-algae symbiotic system, heat-dependent photoinhibition of photosystem II (PSII) leads to coral bleaching. When the reef-building coral Acropora digitifera was exposed to light, a moderate increase of temperature induced coral bleaching through photobleaching of algal pigments, but not through expulsion of symbiotic algae. Monitoring of PSII photoinhibition revealed that heatdependent photoinhibition was ascribed to inhibition of the repair of photodamaged PSII, and heat susceptibility of the repair machinery varied among coral species. We conclude that the efficiency of the photosynthesis repair machinery determines the bleaching susceptibility of coral species under elevated seawater temperatures. Talbot, F and Wilkinson, C. (2001) Coral reefs, mangroves and seagrasses: A sourcebook for managers. Australian Institute of Marine Science: Townsville (Australia), 193 pp. This book is written for resource managers and educators. Its goal is to provide information and practical examples to help prevent further damage to coral reefs and other tropical coastal ecosystems. Already 11% of all coral reefs have been damaged beyond recovery or totally destroyed. Another 16% were wrecked in 1998 by climate change related coral bleaching. Without effective management, another 30% of the world’s reefs will become seriously depleted in the next 20 to 40 years - which is a very short time in human history. While this damage is occurring all over the world, the causes are mostly local – what is happening at each site. This estimate excludes the 1998 coral bleaching caused by higher sea surface temperatures. This bleaching will add to the damage but the full extent is unknown. Most reef damage has been caused by human impacts, usually accidental or unintentional, and is often exaggerated by natural stresses. We now know enough to prevent most of this local damage. More than 50% of the world’s mangrove forests have been cleared without understanding the enormous value of these muddy coastal forests. Seagrass beds have had similar losses in many areas. Both these systems are rich nurseries for coastal fish and prawns, and both act to protect fragile shorelines from erosion. Economists and managers have begun to measure the dollar value of these resources to coastal and marine fisheries, and to shoreline protection. They are finding that these values are surprisingly high; far higher than the money obtained by chopping down mangrove trees or pulling trawl nets through seagrass beds. It makes

good economic sense to manage these coastal resources sustainably and this book will help managers do this. We use real life situations to draw conclusions and give practical advice on the management of coral reefs, mangrove forests and seagrass beds. We outline where there have been real successes and failures, and provide methods for improved management of coastal resources, based on these case studies. Damaged habitats lose much of their economic value and often have bad impacts on other resources nearby. The information in this book is drawn from the best available expertise of scientists and marine resource managers. Our aim is to highlight the dominant issues and suggest achievable solutions to what are sometimes complex situations. We have written the book for the resource manager in the field, who may be working without a lot of scientific support and literature. It is written in basic language and avoids technical terms where possible, because we realise that many managers use English as their second or third language. It will also be useful to fishery managers, regional and town planners, tourist developers, marine park managers, traditional reef owners and others. We include useful references and we have concentrated on material that is available from international agencies like UNESCO, UNEP and IUCN. We particularly refer readers wanting more information to UNESCO's Coral Reef Management Handbook and the IUCN's Guidelines for Marine Protected Areas. Tamelander, J. (2002) Coral recruitment following a mass mortality event. Ambio 31, 551-557. Coral recruitment, following the coral bleaching episode in 1998, was studied on the Kenyan coast. Scleractinian recruits representing 31 genera from 13 families were recorded in 2001, the dominant families at all sites being the Pocilloporidae, Poritidae, and Faviidae. The highest diversity and density of scleractinian recruits was observed at a site located within a marine park, with 11 families, 20 genera and 21.4 recruits per m2, while an unprotected site and sites with higher sediment input showed recruit densities under five. Survivorship was generally higher in massive than in branching genera. The diversity in the recruit population has decreased compared to pre-bleaching levels, while no significant change in density was recorded. Growth rates in recruits were consistent between sites and time of year, with an average growth rate of 0.117 mm2 mm–2 mon–1, with variation between species. The fastest growing genera were Echinopora, Acropora, Pocillopora, and Porites. With the exception of Montipora, the growth rate of surviving pre-bleaching colonies was lower and exhibited greater variability between taxa and sites than among recruits. The Nature Conservancy (2004) R2 – Reef resilience: Building resilience into coral reef conservation, additional tools for managers CDROM Toolkit. The Nature Conservancy: Honolulu (USA). Managers and Practitioners: This toolkit is a first draft that provides you with the latest tools, strategies, and protocols to address coral bleaching and conservation of reef fish spawning aggregations. You are invited to field-test and refine the methods in this draft. In two years, a revised toolkit will be produced based on your recommendations.

This R2—Reef Resilience—toolkit is designed primarily for use by Marine Protected Area (MPA) managers, trainers, and policymakers. Entrusted with protecting coral reefs, they face significant challenges because these ecosystems are highly vulnerable to the demographic, economic, and environmental changes expected during the next century. R2 builds upon the principle that effective management is fundamental to ensure reef survival. Comprised of two extensive topic streams—Coral Bleaching and Reef Fish Spawning Aggregations—the toolkit will help practitioners begin to build resilience into their coral reef conservation programs so that these valuable natural systems can survive the anticipated rapid changes and provide for escalating human needs. This Coral Bleaching Stream examines the definition, causes, and consequences of coral bleaching and recognizes that mass bleaching does not affect all corals equally on any reef system. Building on this, the toolkit provides information and approaches to assist MPA managers examine factors that may help coral communities either to resist or recover quickly from bleaching events. It also helps them identify the locations of resistant communities. These communities provide vital refugia that are essential to the recovery and survival of areas prone to degradation from multiple causes, including mass bleaching. Protection of these refugia is a second underlying principle of the toolkit. The later sections of this toolkit, while constrained by the current state of knowledge on the subject, propose specific actions that MPA managers can take to diminish the impacts of mass bleaching at different scales, from national to local. As our knowledge of the issues surrounding coral bleaching and spawning aggregations improves, and our understanding of better ways to deal with them becomes apparent, new iterations of this toolkit may be anticipated. The second stream of the R2 toolkit focuses on reef fish spawning aggregations as another important source of seed for replenishment of degraded reefs. Companion volume on MPA design and management: http://www.iucn.org/themes/marine/pdf/mpaguid2.pdf The Royal Society (2005) Ocean acidification due to increasing atmospheric carbon dioxide. The Royal Society: London, 60 pp. The oceans cover over two-thirds of the Earth’s surface. They play a vital role in global biogeochemical cycles, contribute enormously to the planet’s biodiversity and provide a livelihood for millions of people. The oceans are absorbing carbon dioxide (CO2) from the atmosphere and this is causing chemical changes by making them more acidic (that is, decreasing the pH of the oceans). In the past 200 years the oceans have absorbed approximately half of the CO2 produced by fossil fuel burning and cement production. Calculations based on measurements of the surface oceans and our knowledge of ocean chemistry indicate that this uptake of CO2 has led to a reduction of the pH of surface seawater of 0.1 units, equivalent to a 30% increase in the concentration of hydrogen ions. If global emissions of CO2 from human activities continue to rise on current trends then the average pH of the oceans could fall by 0.5 units (equivalent to a three fold increase in the concentration of hydrogen ions) by the year 2100. This pH is probably lower than has been experienced for hundreds of millennia and, critically, this rate of change is probably one hundred times greater than at any time over this period. The

scale of the changes may vary regionally, which will affect the magnitude of the biological effects. Ocean acidification is essentially irreversible during our lifetimes. It will take tens of thousands of years for ocean chemistry to return to a condition similar to that occurring at pre-industrial times (about 200 years ago). Our ability to reduce ocean acidification through artificial methods such as the addition of chemicals is unproven. These techniques will at best be effective only at a very local scale, and could also cause damage to the marine environment. Reducing CO2 emissions to the atmosphere appears to be the only practical way to minimise the risk of large-scale and long-term changes to the oceans. All the evidence collected and modelled to date indicates that acidification of the oceans, and the changes in ocean chemistry that accompany it, are being caused by emissions of CO2 into the atmosphere from human activities. The magnitude of ocean acidification can be predicted with a high level of confidence. The impacts of ocean acidification on marine organisms and their ecosystems are much less certain but it is likely that, because of their particular physiological attributes, some organisms will be more affected than others. Predicting the direction and magnitude of changes in a complex and poorly studied system such as the oceans is very difficult. However, there is convincing evidence to suggest that acidification will affect the process of calcification, by which animals such as corals and molluscs make shells and plates from calcium carbonate. The tropical and subtropical corals are expected to be among the worst affected, with implications for the stability and longevity of the reefs that they build and the organisms that depend on them. Cold-water coral reefs are also likely to be adversely affected, before they have been fully explored. Other calcifying organisms that may be affected are components of the phytoplankton and the zooplankton, and are a major food source for fish and other animals. Regional variations in pH will mean that by 2100 the process of calcification may have become extremely difficult for these groups of organisms particularly in the Southern Ocean. Some shallow water animals, which play a vital role in releasing nutrients from sediments, also calcify, and may be affected by changes in the chemistry of the oceans. Some studies suggest that growth and reproduction in some calcifying and non-calcifying marine species could be reduced due to the projected changes in ocean chemistry. From the evidence available it is not certain whether marine species, communities and ecosystems will be able to acclimate or evolve in response to changes in ocean chemistry, or whether ultimately the services that the ocean’s ecosystems provide will be affected. Research into the impacts of high concentrations of CO2 in the oceans is in its infancy and needs to be developed rapidly. We recommend that a major, internationally coordinated effort be launched to include global monitoring, experimental, mesocosm and field studies. Models that include the effects of pH at the scale of the organism and the ecosystem are also necessary. The impacts of ocean acidification are additional to, and may exacerbate, the effects of climate change. For this reason, the necessary funding should be additional and must not be diverted from research into climate change. Oceans play a very important role in the global carbon cycle and Earth’s climate system. There are potentially important interactions and feedbacks between changes in the state of the oceans (including their pH) and changes in the global climate and atmospheric chemistry. Changes in the chemistry of the oceans will reduce their ability to absorb additional CO2 from the atmosphere, which will in turn affect the rate

and scale of global warming. The knowledge of these impacts and effects is currently poor and requires urgent consideration. The understanding of ocean acidification and its impacts needs to be taken into account by the Intergovernmental Panel on Climate Change and kept under review by international scientific bodies such as the Intergovernmental Oceanographic Commission, the Scientific Committee on Oceanic Research and the International Geosphere-Biosphere Programme. The socio-economic effects of ocean acidification could be substantial. Damage to coral reef ecosystems and the fisheries and recreation industries that depend on them could amount to economic losses of many billions of dollars per year. In the longer term, changes to the stability of coastal reefs may reduce the protection they offer to coasts. There may also be direct and indirect effects on commercially important fish and shellfish species. Marine ecosystems are likely to become less robust as a result of the changes to the ocean chemistry and these will be more vulnerable to other environmental impacts (such as climate change, water quality, coastal deforestation, fisheries and pollution). The increased fragility and sensitivity of marine ecosystems needs to be taken into consideration during the development of any policies that relate to their conservation, sustainable use and exploitation, or the communities that depend on them. If the risk of irreversible damage arising from ocean acidification is to be avoided, particularly to the Southern Ocean, the cumulative future human derived emissions of CO2 to the atmosphere must be considerably less than 900 Gt C (gigatonnes of carbon) by 2100. In setting targets for reductions in CO2 emissions, world leaders should take account of the impact of CO2 on ocean chemistry, as well as on climate change. These targets must be informed by sound science. Ocean acidification is a powerful reason, in addition to that of climate change, for reducing global CO2 emissions. Action needs to be taken now to reduce global emissions of CO2 to the atmosphere to avoid the risk of irreversible damage to the oceans. We recommend that all possible approaches be considered to prevent CO2 reaching the atmosphere. No option that can make a significant contribution should be dismissed. Thornhill, DJ, LaJeunesse, TC, Kemp, DW, Fitt, WK and Schmidt, GW. (2006) Multi-year, seasonal genotypic surveys of coral-algal symbioses reveal prevalent stability or post-bleaching reversion. Marine Biology 148, 711-722. This report documents the extent to which coral colonies show fluctuations in their associations with different endosymbiotic dinoflagellates. The genetic identity of Symbiodinium from six coral species [Acropora palmata (Lamarck), A. cervicornis (Lamarck), Siderastrea siderea (Ellis and Solander), Montastrea faveolata (Ellis and Solander), M. annularis (Ellis and Solander), and M. franksi (Gregory)] was examined seasonally over five years (1998 and 2000–2004) in the Bahamas and Florida Keys at shallow (1 to 4 m) forereef/patch reef sites and at deeper fore-reef (12–15 m) locations. Symbionts were identified genetically using denaturing gradient gel electrophoresis (DGGE) fingerprinting of the internal transcribed spacer region 2 (ITS2) of ribosomal RNA gene loci. Repetitive sampling from most labeled colonies from the Bahamas and the Florida Keys showed little to no change in their dominant symbiont. In contrast, certain colonies of M. annularis and M. franksi from the Florida Keys exhibited shifts in their associations attributed to recovery from the stresses of the 1997–1998 El Niño southern oscillation (ENSO) event. Over several years, a

putatively stress-tolerant clade D type of Symbiodinium was progressively replaced in these colonies by symbionts typically found in M. annularis and M. franksi in Florida and at other Caribbean locations. Greater environmental fluctuations in Florida may explain the observed changes among some of the symbioses. Furthermore, symbiotic associations were more heterogeneous at shallow sites, relative to deep sites. The exposure to greater environmental variability near the surface may explain the higher symbiont diversity found within and between host colonies. Tilmant, J (ed.) (2000) Coral reef protected areas: A guide for management. US Coral Reef Taskforce: Washington DC (USA), 14 pp. Abstract not available. Tioho, H, Tokeshi, M, Nojima, S. (2001) Experimental analysis of recruitment in a scleractinian coral at high latitude. Marine Ecology Progress Series 213, 79-86. A set of field experiments were conducted to determine the patterns of dispersal and recruitment in the brooding scleractinian coral Pocillopora damicornis at high-latitude (32°N), temperate locations in Amakusa, south-western Japan. Planulation occurred only in July (from full moon to last-quarter moon) in 2 consecutive years, confirming a pattern of annual reproduction in this species at the study site. Settlement plates placed beside naturally occurring colonies of P. damicornis accumulated more recruits than plates placed away (8 m) from the colonies in both years. Another experiment involving transplantation of P. damicornis colonies into an area where the species did not naturally occur but where environmental conditions were considered adequate for its growth also demonstrated that the density of settling planulae was high close to parent colonies but declined steadily with increasing distance. Planulae distribution tended to be more aggregated with increasing distance from their source. The present study demonstrates that the planulae of brooding coral species do not generally disperse over long distances at high latitude. Tkachenko, KS, Wu, B-J, Fang, L-S and Fan, T-Y. (2007) Dynamics of a coral reef community after mass mortality of branching Acropora corals and an outbreak of anemones. Marine Biology 151, 185–194. Dynamics of a coral reef community at Tiao-Shi Reef, southern Taiwan were studied using permanent transects to examine coral recovery and successive cascades to collapse stage resulting from chronic anthropogenic impacts and typhoons. Three distinct zones were recognized within a relatively small study area (250 m across) formerly dominated by large stands of branching Acropora corals. The first zone still retains the dominance of branching Acropora corals, although they show a significant decreasing tendency. The second zone exhibits recovery with a significant increase in branching Montipora stellata, which is recruited and grows faster than branching Acropora corals. The third zone is occupied by anemone, Condylactis sp., and demonstrates a stable phase of coral deterioration without recovery. Such differences in coral reef community dynamics within a small spatial scale illustrate mosaic dynamics which have resulted from degradation of the water quality, patchy mortality of large branching Acropora thickets caused by typhoons, the rapid asexual fragmentation and growth of M. stellata making it a successful colonizer, and

occupation by anemone, Condylactis sp., together with unstable remnants of dead Acropora rubbles have not allowed coral recruits to survive. Toller, WW, Rowan, R and Knowlton, N. (2001) Repopulation of zooxanthellae in the Caribbean corals Montastrea annularis and M. faveolata following experimental and disease-associated bleaching. The Biological Bulletin 201, 360-373. Caribbean corals of the Montastraea annularis species complex associate with four taxa of symbiotic dinoflagellates (zooxanthellae; genus Symbiodinium) in ecologically predictable patterns. To investigate the resilience of these host-zooxanthella associations, we conducted field experiments in which we experimentally reduced the numbers of zooxanthellae (by transplanting to shallow water or by shading) and then allowed treated corals to recover. When depletion was not extreme, recovering corals generally contained the same types of zooxanthellae as they did prior to treatment. After severe depletion, however, recovering corals were always repopulated by zooxanthellae atypical for their habitat (and in some cases atypical for the coral species). These unusual zooxanthellar associations were often (but not always) established in experimentally bleached tissues even when adjacent tissues were untreated. Atypical zooxanthellae were also observed in bleached tissues of unmanipulated Montastraea with yellow-blotch disease. In colonies where unusual associations were established, the original taxa of zooxanthellae were not detected even 9 months after the end of treatment. These observations suggest that zooxanthellae in Montastraea range from fugitive opportunists and stress-tolerant generalists (Symbiodinium A and E) to narrowly adapted specialists (Symbiodinium B and C), and may undergo succession. Toscano, MA, Liu, G, Guch, IC, Casey, KS, Strong, AE and Meyer, JE. (2000) Improved prediction of coral bleaching using high-resolution HotSpot anomaly mapping. 9th International Coral Reef Symposium: Bali (Indonesia), 6 pp. NOAA/NESDIS developed global, 50km satellite SST HotSpot anomaly maps to provide early warnings of thermally-induced coral bleaching. Hotspots are anomalies above a satellite-derived climatological Maximum Monthly Mean (MMM) SST. An increase to 1°C above the MMM SST during summer is a general threshold for inducing coral bleaching. 50km HotSpot resolution predicted bleaching over broad areas during a major climatic event (1998 ENSO); however, regional and reef-scale (meters to tens of kilometers) warming events may induce coral bleaching. To test reef-scale monitoring, a 9km retrospective (1998) HotSpot mapping study used NASA/JPL AVHRR Oceans Pathfinder Best SST data. Pathfinder SSTs accurately reproduced in situ temperatures, and were used to create a 9km MMM threshold climatology for highresolution HotSpot mapping. Warmer 9km climatology values throughout the tropics generally reduced anomaly levels from the 50km product, suggesting that coral bleaching may be triggered by minimal thermal stress.

Toscano, MA, Casey, KS and Shannon, J. (2002) Use of high resolution Pathfinder SST data to document coral reef bleaching. Seventh International Conference on Remote Sensing for Marine and Coastal Environments, Miami (Florida), 8 pp. Mass coral bleaching has been observed throughout the tropical oceans immediately following periods of thermal stress, in concert with other known causes. While water temperatures monitored in situ best document the relationship of thermal stress to bleaching, most reef sites have limited to no in situ data. For such areas, high-resolution satellite-derived SSTs provide valuable time series data and quantify thermal anomalies and their timing vs. onset of bleaching. The availability of 9km NOAA/NASA Oceans Pathfinder SST data (1985-2001), an internally consistent, calibrated SST database at reef scale resolution, allows us to assess the accuracy and usefulness of satellite SST data to document past conditions in reefs. Statistical analyses of SSTs combined with available in situ temperatures from Caribbean and tropical Pacific locations indicate that 9km SSTs for specific sites, and for the 3x3 pixel means surrounding each site, are highly correlated to in situ temperatures. Biases are tied, wherever possible, to local physical causes. Pathfinder SSTs accurately represent surface field temperatures with predictable exceptions, and therefore have great potential for assessing temperature histories of remote, un-monitored reefs Underwood, JN, Smith, LD and van Oppen, MJH. (2007) Multiple scales of genetic connectivity in a brooding coral on isolated reefs following catastrophic bleaching. Molecular Ecology 16, 771–784. Blackwell Publishing Ltd Understanding the pattern of connectivity among populations is crucial for the development of realistic and spatially explicit population models in marine systems. Here we analysed variation at eight microsatellite loci to assess the genetic structure and to infer patterns of larval dispersal for a brooding coral, Seriatopora hystrix, at an isolated system of reefs in northern Western Australia. Spatial autocorrelation analyses show that populations are locally subdivided, and that the majority of larvae recruit to within 100 m of their natal colony. Further, a combination of F- and R- statistics showed significant differentiation at larger spatial scales (2–60 km) between sites, and this pattern was clearly not associated with distance. However, Bayesian analysis demonstrated that recruitment has been supplemented by less frequent but recent input of larvae from outside the local area; 2–6% of colonies were excluded from the site at which they were sampled. Individual assignments of these migrants to the most likely populations suggest that the majority of migrants were produced at the only site that was not decimated by a recent and catastrophic coral bleaching event. Furthermore, the only site that recovered to prebleaching levels received most of these immigrants. We conclude that the genetic structure of this brooding coral reflects its highly opportunistic life history, in which prolific, philopatric recruitment is occasionally supplemented by exogenously produced larvae.

United States Coral Reef Task Force (2000) The national action plan to conserve coral reefs. USCRTF: Washington DC (USA), 34 pp. Coral reefs are among the most diverse and biologically complex ecosystems on earth. These rainforests of the sea provide economic and environmental services to millions of people as areas of natural beauty and recreation, sources of food, jobs, chemicals, pharmaceuticals, and shoreline protection. Now under threat from multiple stresses that are overwhelming their natural resilience, coral reefs are deteriorating worldwide at alarming rates. An estimated 10% of the world’s reefs have already been lost and 60% are threatened by bleaching, disease and a variety of human activities including shoreline development, polluted runoff from agricultural and land-use practices, ship groundings, over-harvesting, destructive fishing, and global climate change. The trend in coral reef health is downward, and these ancient ecosystems are in peril. This document presents the United States’ collective response to this crisis: The National Action Plan to Conserve Coral Reefs. The Action Plan was produced by the Working Groups of the United States Coral Reef Task Force in response to its request for a cohesive national strategy to implement Executive Order 13089 on Coral Reefs (Appendix A). These actions were developed in consultation with a variety of stakeholders and cover the spectrum of coral reef conservation from mapping, monitoring, management and research, to education and international cooperation (Appendix B). Collectively, these actions are intended to provide a comprehensive road map for federal, state, territorial and local action to reverse the worldwide decline and loss of coral reefs. This is a living document, intended by its authors to be revisited and revised regularly, and to be augmented by agency implementation plans and an annual report from each CRTF member agency summarizing significant issues and accomplishments related to coral reef conservation. Strategies to Conserve Coral Reefs Two fundamental themes will frame the Coral Reef Task Force’s conservation actions: Ø Understand Coral Reef Ecosystems – by conducting comprehensive mapping, assessment and monitoring of coral reef habitats; supporting strategic research on regional threats to coral reef health and the underlying ecological processes upon which they depend; and incorporating the human dimension into conservation and management strategies. Ø Reduce The Adverse Impacts Of Human Activities – by creating an expanded and strengthened network of federal, state and territorial coral reef Marine Protected Areas; reducing the adverse impacts of extractive uses; reducing habitat destruction; reducing pollution; restoring damaged reefs; strengthening international activities; reducing the impacts of international trade in coral reef species; improving governmental accountability and coordination; and creating an informed and engaged public for coral reef conservation. In addition to these specific action strategies, the Coral Reef Task Force member agencies have developed a suite of 8 core principles to guide their future endeavors to implement the Executive Order as well as the National Action Plan: Ø to adopt a science-based ecosystem approach to coral reef conservation that recognizes and builds upon important linkages among adjacent and remote habitats associated with coral reefs.

Ø to employ adaptive management approaches that track and respond to environmental change and emerging threats. Ø that scientific uncertainty shall not prevent taking precautionary measures as appropriate to protect coral reefs. Ø to incorporate the human dimension into coral reef conservation strategies by ensuring that management measures reflect, and are sensitive to the local socioeconomic, political and cultural environment, and that they build an informed public engaged in choosing alternatives to activities that harm coral reefs. Ø to apply marine zoning - including marine protected areas and no-take ecological reserves - in order to protect and replenish coral reef ecosystems by minimizing harmful human impacts and user conflicts in important habitats. Ø to fully and proactively use existing management authorities and programs at the federal, state and territorial levels, and develop, where needed, new legal mechanisms that protect, restore and enhance coral reef ecosystems. Ø to develop and support strong domestic partnerships among governmental, private and scientific interests to meet the complex cross-jurisdictional challenges of coral reef conservation. Ø to provide global leadership to reduce global threats to coral reefs through international technical and development assistance, capacity building and collaboration. Valentine, JF and Heck Jr, KL. (2005) Perspective review of the impacts of overfishing on coral reef food web linkages. Coral Reefs 24, 209-213. Overfishing is increasingly understood to result in indirect alterations of habitat structure and function (McClanahan et al. 1995; Bowen 1997; Jackson 2001; Jackson et al. 2001; Gardner et al. 2003). The intense harvesting of sea otters from kelp forests, for example, created ‘‘low- predation refuges’’ for their sea urchin prey. Following otter removal, sea urchin grazing exploded resulting in large regional losses of kelp (Estes and Palmisano 1974; Duggins 1980). Further support for the impacts of large consumers was found in Alaskan embayments where inshore migration of killer whales changed the strength of interactions between sea otters and their herbivorous sea urchin prey, again resulting in local losses of kelp forests (Estes et al. 1998). Similarly, McClanahan et al. (1995) found that intense harvesting of predatory trigger fishes promoted large-scale habitat changes as their sea urchin prey, once released from predation, increased. More urchins subsequently led to more coral reef erosion and the eventual replacement of corals by seagrasses (McClanahan and Kurtis 1991). Moreover, the intense harvesting of both large piscivorous and herbivorous fishes, coupled with the die-off of herbivorous sea urchins, is thought to be responsible for widespread macroalgal overgrowth of coral reefs in Jamaica (Hughes 1994 but see Aronson and Precht 2001). Given that the dramatic impacts of fishing on marine food webs preceded the development of most theory in marine ecology, it is understandable that management agencies have predominantly focused their efforts on the role of bottom-up processes (nutrient concentration) in regulating the productivity of coastal ecosystems. But given the increasing evidence that food web alterations have figured prominently in the collapse of marine ecosystems, it seems clear that new studies are urgently needed to assess the degree to which large-scale food web alterations have changed the structure and function of marine ecosystems (Estes and Peterson 2000; Jackson 2001; Pandolfi et al. 2003).

Van Oppen, MJH, Mahiny, AJ and Done, TJ. (2005) Geographic distribution of zooxanthellae types in three coral species on the Great Barrier Reef sampled after the 2002 bleaching event. Coral Reefs 24, 482-487. Reef corals form obligate symbiotic relationships with many genetically divergent taxa of zooxanthellae (dinoflagellates of the genus Symbiodinium) (reviewed in Baker 2003). Based on nuclear ribosomal DNA (rDNA) and later confirmed by partial chloroplast large subunit rDNA sequences (Santos et al. 2002), eight distinct phylogenetic clades of zooxanthellae have been distinguished (A-H; LaJeunesse 2001; Rodriguez-Lanetty 2003; Pochon et al. 2004), five of which (A-D, F) are known to form associations with scleractinian corals. This high taxon diversity is accompanied by differences in physiological traits between some zooxanthella types (Kinzie et al. 2001; Bhagooli et al. 2003; Little et al. 2004), with Symbiodinium clade D being the most heat-tolerant type known today (Glynn et al. 2001; Baker et al. 2004; Fabricius et al. 2004; Rowan 2004). Physiological differences among zooxanthella types has led to the expectation that zooxanthella types may affect their hosts’ sensitivity to environmental conditions and/or that changes in environmental conditions may affect the distribution of zooxanthella types. In January and February 2002, the Great Barrier Reef (GBR) was exposed to an extended period of elevated sea water temperature and high light that led to widespread but patchy coral bleaching (expulsion of zooxanthellae by the coral host and/or loss of zooxanthellar pigments) (Berkelmans et al. 2004). To search for possible links between the zooxanthella type harbored by the coral colonies and the extent of damage suffered from the bleaching event, and also to explore patterns of zooxanthella distribution, we sampled zooxanthellae from three common species of scleractinian corals, Acropora millepora, A. tenuis and Stylophora pistillata, from 17 sites along a latitudinal and cross-shelf gradient in the central and southern sections of the GBR 4–5 months after the 2002 bleaching event. The health status of each coral colony was recorded at the time of collection, and its zooxanthella genotype identified, using single stranded conformation polymorphism (SSCP) and sequence analysis of the nuclear ribosomal DNA ITS1 region. There were no simple correlations between symbiont types and either the level of bleaching of individual colonies or indicators of heat stress at individual sites. However, there was a very high post-bleaching abundance of the heat tolerant symbiont type D (Baker et al. 2004; Fabricius et al. 2004; Rowan 2004) in one coral population at the most heat-stressed site. Van Oppen, MJH, Mieog, JC, Sánchez, CA and Fabricius, KE. (2005) Diversity of algal endosymbionts (zooxanthellae) in octocorals: the roles of geography and host relationships. Molecular Ecology 14, 2403–2417. The presence, genetic identity and diversity of algal endosymbionts (Symbiodinium) in 114 species from 69 genera (20 families) of octocorals from the Great Barrier Reef (GBR), the far eastern Pacific (EP) and the Caribbean was examined, and patterns of the octocoral–algal symbiosis were compared with patterns in the host phylogeny. Genetic analyses of the zooxanthellae were based on ribosomal DNA internal transcribed spacer 1 (ITS1) region. In the GBR samples, Symbiodinium clades A and G were encountered with A and G being rare. Clade B zooxanthellae have been previously reported from a GBR octocoral, but are also rare in octocorals from this region. Symbiodinium G has so far only been found in Foraminifera, but is rare in

these organisms. In the Caribbean samples, only Symbiodinium clades B and C are present. Hence, Symbiodinium diversity at the level of phylogenetic clades is lower in octocorals from the Caribbean compared to those from the GBR. However, an unprecedented level of ITS1 diversity was observed within individual colonies of some Caribbean gorgonians, implying either that these simultaneously harbour multiple strains of clade B zooxanthellae, or that ITS1 heterogeneity exists within the genomes of some zooxanthellae. Intracladal diversity based on ITS should therefore be interpreted with caution, especially in cases where no independent evidence exists to support distinctiveness, such as ecological distribution or physiological characteristics. All samples from EP are azooxanthellate. Three unrelated GBR taxa that are described in the literature as azooxanthellate (Junceella fragilis, Euplexaura nuttingi and Stereonephthya sp. 1) contain clade G zooxanthellae, and their symbiotic association with zooxanthellae was confirmed by histology. These corals are pale in colour, whereas related azooxanthellate species are brightly coloured. The evolutionary loss or gain of zooxanthellae may have altered the light sensitivity of the host tissues, requiring the animals to adopt or reduce pigmentation. Finally, we superimposed patterns of the octocoral–algal symbiosis onto a molecular phylogeny of the host. The data show that many losses/gains of endosymbiosis have occurred during the evolution of octocorals. The ancestral state (azooxanthellate or zooxanthellate) in octocorals remains unclear, but the data suggest that on an evolutionary timescale octocorals can switch more easily between mixotrophy and heterotrophy compared to scleractinian corals, which coincides with a low reliance on photosynthetic carbon gain in the former group of organisms. Van Oppen, MJ and Gates, RD. (2006) Conservation genetics and the resilience of reef-building corals. Molecular Ecology 15, 3863-3883. Coral reefs have suffered long-term decline due to a range of anthropogenic disturbances and are now also under threat from climate change. For appropriate management of these vulnerable and valuable ecosystems it is important to understand the factors and processes that determine their resilience and that of the organisms inhabiting them, as well as those that have led to existing patterns of coral reef biodiversity. The scleractinian (stony) corals deposit the structural framework that supports and promotes the maintenance of biological diversity and complexity of coral reefs, and as such, are major components of these ecosystems. The success of reef-building corals is related to their obligate symbiotic association with dinoflagellates of the genus Symbiodinium. These one-celled algal symbionts (zooxanthellae) live in the endodermal tissues of their coral host, provide most of the host's energy budget and promote rapid calcification. Furthermore, zooxanthellae are the main primary producers on coral reefs due to the oligotrophic nature of the surrounding waters. In this review paper, we summarize and critically evaluate studies that have employed genetics and/or molecular biology in examining questions relating to the evolution and ecology of reef-building corals and their algal endosymbionts, and that bear relevance to coral reef conservation. We discuss how these studies can focus future efforts, and examine how these approaches enhance our understanding of the resilience of reef-building corals.

Van Rooij, JM, Videler, JJ and Bruggemann, JH. (1998) High biomass and production but low energy transfer efficiency of Caribbean parrotfish: Implications for trophic models of coral reefs. Journal of Fish Biology 53, 154-178. Quantitative data are presented to assess the trophic role of scarids on the fringing coral reef of Bonaire (Netherlands Antilles), with particular emphasis on the stoplight parrotfish Sparisoma viride. Average herbivore biomass on the reef was 690 kg ha-1, 22% of which was accounted for by S. viride. From data on relative gonad weights, daily spawning frequencies, and egg numbers obtained by stripping, with previous estimates of somatic growth and energy intake, a gross efficiency (GE: somatic plus gamete production/ consumption) of 2·3% was obtained. This is a factor of five to seven lower than the GE suggested to be valid for most aquatic ecosystems, including coral reefs. To investigate one potential cause for our low estimate, overestimation of food intake, our intake estimates were compared with published values for other herbivorous coral reef fish. This yielded a relationship (daily C intake=0·0342xW0·816; wet body mass W in g) with high correlation (r2=94·6%, n=13), which shows that the intake estimates agree well with other published data. Averaged over the year, primary production at 0–3 m depth was 17·2 kg C ha-1 day-1 while herbivore consumption was estimated at 17·4 kg C ha-1 day-1, indicating an ecotrophic efficiency (EE, the fraction of total production at one trophic level that is consumed by all predators) of 100%. This suggests strongly that the food intake estimates are realistic, since no changes in algal biomass were observed over the study period. The two scarids for which food intake was actually measured in our own study area, were estimated to consume 55% of the algal production in the shallow reef (S. viride, 20%; Scarus vetula, 35%). This is lower than expected if consumption were proportional to biomass (S. viride, 22%; S. vetula, 40% of herbivore biomass in the shallow reef). Consequently, a minimum estimate of 88–91% can be inferred for the EE of these two species. Multiplied by GE, this yields a transfer efficiency (TE, the fraction of production passing from one trophic level to the next) of 2%. For coastal and coral systems the primary production required (PPR) to sustain fisheries was estimated to be 8·3%, which was based on a TE of 10%. The present estimates show that the TE of a major herbivore at our reef is at least a factor of five lower. Assuming that the estimate is representative for all scarids (comprising 70% of the herbivore standing stock), it can be concluded that the PPR to sustain coral reef fisheries may be as high as 40% of the total primary production. The low value reported before, might suggest that the effect of fishing mainly affects target populations but not the lowest trophic levels. It is argued that our estimate is more realistic for coral reefs supporting high scarid biomass and explains better the many reports of coral destruction due to algal overgrowth at exploited reefs. Van Woesik, R. (2001) Coral bleaching: Transcending spatial and temporal scales. Trends in Ecology and Evolution 16, 119-121. Corals on tropical coral reefs maintain their colourful, almost surrealistic appearance throughout the year, unless they ‘bleach’ when the waters get too warm. The bleaching of corals through loss of pigmentation or symbiotic algae (zooxanthellae) can be temporary or can lead to coral mortality. At the recent International Coral Reef Symposium, coral bleaching was the subject of a mini symposium, where, among

other topics, the 1997–1998 coral-bleaching event and associated coral mortality, reported from over 50 countries, was discussed. Van Woesik, R. (2002) Processes regulating coral communities. Comments on Theoretical Biology 7, 201-214. While coral reefs support more species than any other marine system, coral reef ecology borrows heavily from theoretical advances in low-diversity terrestrial systems. This article reviews recent developments in ecology on coral reefs and weaves together the relevant information within a theoretical framework that aims toward predictive modeling of coral populations and communities. The author begins with a historical perspective, involving arguments that shaped modern community theory, and introduces coral communities within this context. The major parameters that shape coral community structure are outlined and the question of whether competition between corals and disturbance frequency actually drives diversity and local community structure i.f discussed. Within the context of metapopulation modeling, involving colonization and extinction events, the author discusses reef connectivity and postsettlement processes and argues for a neutral model with high diversity as the default system. The article concludes with a brief discussion on deficiencies in coral community data and future modeling directions. Vermeij, MJA. (2006) Early life-history dynamics of Caribbean coral species on artificial substratum: the importance of competition, growth and variation in life-history strategy. Coral Reefs 25, 59–71. The development of a coral community was monitored for 6 years (1998–2004) on 46 m2 of artificial settlement substrate in Curaçao, Netherlands Antilles. Growth and survival of recruits (n=1385) belonging to 16 different species were quantified in relation to characteristics of the benthic community developing around them. The early life history dynamics (i.e. growth rate, growth strategy and survival) of corals differed among species although these differences were small for species occupying similar habitats (i.e. underside versus topside of substratum). In contrast to recruit survival, juvenile growth rates were highly variable and unrelated to benthic community structure, at least at the scale of this study. Competing benthic organisms affected coral recruitment success through space preemption (mainly by macroalgae) or recruit overgrowth (mainly by sponges). The results highlight the small spatial scale (mm–cm) at which the processes responsible for recruitment success or failure occur and emphasize the need to include such small-scale observations in studies of coral early life-phase dynamics. Vestergaard, O, Hoegh-Guldberg, O and Unluata, U. (2003) Understanding coral bleaching across four oceans: Addressing CBD’s specific workplan on coral bleaching. Convention on Biological Diversity: Montreal (Canada), 6 pp. Coral bleaching is rapidly developing as a major problem for the health of coral reefs worldwide. Unfortunately, the level of understanding of this phenomenon is limited. Developing a good understanding of the phenomenon is important if management practices are to be effective in minimising the detrimental impacts of coral bleaching as projected to occur over the next 50 years.

In response to this situation an expert group, the IOC/UNESCO Working Group on Coral Bleaching, has been established with the purpose to 1) integrate, synthesize and develop global research on coral bleaching and related impacts of climate change on coral ecosystems; 2) generate tools and techniques to detect early stress responses and longer-term impacts of climate change with a wide applicability in both developed and developing nations; and 3) strengthen the capacity for research and management of coral bleaching impacts in developing countries affected by coral bleaching, through the transfer of expertise and technologies. The effort has become an important implementer of CBD’s Specific Workplan on Coral Bleaching (COP 6 VI/3, 2002). Visram, S and Douglas, AE. (2006) Molecular diversity of symbiotic algae (zooxanthellae) in scleractinian corals of Kenya. Coral Reefs 25, 172-176. In this first sequence analysis of ‘zooxanthellae’ (symbiotic algae of the genus Symbiodinium) in scleractinian corals in Africa, seven Kenyan species sampled in 2001–2002 were analysed by RFLP and sequencing of a PCR-amplified fragment of the LSU rRNA gene. Zooxanthellae of phylotypes A, C and D, all described previously in corals from other regions of the world, were detected. All sequences of phylotype D were identical, while phylotype C was variable, with 14 distinct sequences, seven of which clustered in a previously unreported subgroup of phylotype C, among the 22 samples. These data on the diversity of zooxanthellae in Kenyan corals 3–4 years after the 1998 bleaching event are of potential value for longitudinal studies of temporal changes in zooxanthella diversity in Kenyan corals, especially in relation to future large-scale bleaching episodes. Visram, S and Douglas, AE. (2007) Resilience and acclimation to bleaching stressors in the scleractinian coral Porites cylindrica. Journal of Experimental Marine Biology and Ecology 349, 35–44. ‘Resilience’, the capacity of the coral symbiosis with dinoflagellate algal symbionts (‘zooxanthellae’) to recover after bleaching, is a little-studied but crucial aspect of coral responses to bleaching stressors. This study investigated the response of the zooxanthella population in the coral Porites cylindrica after bleaching either naturally on a shallow subtidal reef or experimentally in response to elevated temperature and darkness. Coral resilience was influenced by the nature and duration of the stressor. Corals strongly bleached by natural stressors were less resilient than those that had been partially bleached; and a similar recovery profile was obtained for corals experimentally bleached by exposure to elevated temperature, in which recovery was slower for corals thermally-stressed 96 h than for 72 h. The opposite trend was evident for corals exposed to darkness, indicating that the bleaching trigger had a strong impact on coral resilience. When P. cylindrica recently recovered from bleaching was subjected to a repetition of bleaching stressors, it did not display acclimation, i.e. experience-mediated acquisition of resistance to bleaching stressors. The zooxanthella populations in all corals tested throughout the experiments were typed by PCR-RFLP as clade C, indicating that coral responses were not accompanied by any substantial change in zooxanthella composition at the cladal level.

Wakeford, M, Done, TJ and Johnson, CR. (2007) Decadal trends in a coral community and evidence of changed disturbance regime. Coral Reefs. A 23 year data set (1981–2003 inclusive) and the spatially explicit individual-based model ‘‘Compete©’’ were used to investigate the implications of changing disturbance frequency on cover and taxonomic composition of a shallow coral community at Lizard Island, Australia. Near-vertical in situ stereo-photography was used to estimate rates of coral growth, mortality, recruitment and outcomes of pair-wise competitive interactions for 17 physiognomic groups of hard and soft corals. These data were used to parameterise the model, and to quantify impacts of three acute disturbance events that caused significant coral mortality: 1982—a combination of coral bleaching and Crown-of-Thorns starfish; 1990—cyclone waves; and 1996—Crown-of-Thorns starfish. Predicted coral community trajectories were not sensitive to the outcomes of competitive interactions (probably because average coral cover was only 32% and there was strong vertical separation among established corals) or to major changes in recruitment rates. The model trajectory of coral cover matched the observed trajectory accurately until the 1996 disturbance, but only if all coral mortality was confined to the 3 years of acute disturbance. Beyond that date (1997–2003), when the observed community failed to recover, it was necessary to introduce annual chronic background mortality to obtain a good match between modelled and observed coral cover. This qualitative switch in the model may reflect actual loss of resilience in the real community. Simulated over a century, an 8 year disturbance frequency most closely reproduced the mean community composition observed in the field prior to major disturbance events. Shorter intervals between disturbances led to reduced presence of the dominant hard coral groups, and a gradual increase in the slow growing, more resilient soft corals, while longer intervals (up to 16 years) resulted in monopolization by the fastest growing table coral, Acropora hyacinthus. Walker, B, Carpenter, S, Anderies, J, Abel, N, Cumming, G, Janssen, M, Lebel, L, Norberg, J, Peterson, GD and Pritchard, D. (2002) Resilience management in social-ecological systems: A working hypothesis for a participatory approach. Conservation Ecology 6. Approaches to natural resource management are often based on a presumed ability to predict probabilistic responses to management and external drivers such as climate. They also tend to assume that the manager is outside the system being managed. However, where the objectives include long-term sustainability, linked social-ecological systems (SESs) behave as complex adaptive systems, with the managers as integral components of the system. Moreover, uncertainties are large and it may be difficult to reduce them as fast as the system changes. Sustainability involves maintaining the functionality of a system when it is perturbed, or maintaining the elements needed to renew or reorganize if a large perturbation radically alters structure and function. The ability to do this is termed “resilience.” This paper presents an evolving approach to analyzing resilience in SESs, as a basis for managing resilience. We propose a framework with four steps, involving close involvement of SES stakeholders. It begins with a stakeholder-led development of a conceptual model of the system, including its historical profile (how it got to be what it is) and preliminary assessments of the drivers of the supply of key ecosystem goods and services. Step 2 deals with identifying the range of unpredictable and uncontrollable drivers, stakeholder visions for the future, and contrasting possible

future policies, weaving these three factors into a limited set of future scenarios. Step 3 uses the outputs from steps 1 and 2 to explore the SES for resilience in an iterative way. It generally includes the development of simple models of the system’s dynamics for exploring attributes that affect resilience. Step 4 is a stakeholder evaluation of the process and outcomes in terms of policy and management implications. This approach to resilience analysis is illustrated using two stylized examples. Walker, D, Solandt, J-L, Haycock, S, Taylor, J, Harding, S and Raines, P. (2002) Fiji coral reef conservation project: Reef Check report (2001-2002). Coral Cay Conservation: London (UK), 27 pp. Reef Check surveys were carried out by CCC volunteer divers in the Mamanucas region, Fiji at the same locations over a two year period (2001-2002) as part of the ‘Fiji Reef Conservation Project’ (FRCP). · Data was collected on anthropogenic impacts, benthic cover and a number of target reef fish and invertebrates at 22 sites. · The results for benthic cover have been reported and discussed with reference to two coral bleaching events. The first, in early 2000 was severe and resulted in significant loss of hard coral cover in the Mamanucas region. The second smaller event occurred in April 2002. · Data reported here indicates that mean hard coral cover has almost doubled between the two survey periods from 14 % in 2001 to 27 % in 2002. There was a corresponding decrease in the amount of reef rock recorded by divers. · Benthic data suggests that the reefs of the Mamanucas region are recovering well after the deleterious effect of coral bleaching in 2000. The smaller bleaching event of 2002 has not significantly altered the amount of live hard coral in the region. · Data collected on target reef fish and invertebrates has highlighted the apparent rarity of some species in the Mamanucas although further information is required to confirm their scarcity. Crown-of-thorns sea stars (COTs) were recorded but at low levels. · Future CCC Reef Check surveys in the Mamanucas (2003 and beyond) will provide evidence to determine whether the data presented here is part of an ongoing trend in terms of coral recovery. Wallace, CC, Muir, PR and Venkatesh, M. (2007) Post-bleaching renewal of the dominant reef-building coral Acropora abrotanoides in the Lakshadweep islands of India. Coral Reefs 26, 45. In the Lakshadweep Islands, SW India, more than 81% of corals died following the 1998 bleaching: rapid but spatially variable onset of recovery followed, with the genus Acropora dominating early recruitment. A survey of Acropora diversity in February 2006 recorded 35 species at 22 sites on Agatti, Kavaratti and Bangaram. Sites were dominated by recovering assemblages of 13–21 Acropora species, with 19 common species occurring in 50–80% of all sites.

Ward, S, Harrison, P and Hoegh-Guldberg, O. (2002) Coral bleaching reduces reproduction of scleractinian corals and increases susceptibility to future stress. Proceedings of the 9th International Symposium for Reef Studies: Bali (Indonesia), 1123-1128. Extensive bleaching of corals occurred at Heron Island Reef during the 1998 mass bleaching event on the Great Barrier Reef, Australia. Two hundred coral colonies were labelled and sampled on the reef flat at Heron Island in March 1998 (when approximately 80% of corals growing in the intertidal reef flat were bleached). These included both bleached and unbleached colonies of many species. Bleached colonies of all sampled species had lower densities of symbiotic dinoflagellates and lower chlorophyll a concentrations per surface area of coral tissue than unbleached colonies. Samples of the colonies were decalcified and polyps dissected to determine fecundity. There were significantly fewer eggs present in the bleached than unbleached colonies in all sampled species. In many species, no eggs were present in the bleached colonies. Eggs that were present were also significantly smaller in the bleached than the unbleached colonies of the majority of species sampled. There were also significantly fewer polyps containing eggs and testes in the bleached than the unbleached colonies. The percentage of tissue made up by lipids in the bleached colonies was significantly lower than that of the unbleached colonies in some species. By July 1998, 23% of the sampled colonies had died and many of the previously bleached colonies had regained their colour, suggesting visually that they had recovered. However, previously bleached colonies in November 1998 still had fewer eggs and reproductive polyps than colonies that had been previously unbleached. In July of the following year, in the middle of the Australian winter, many of the corals that had bleached the year before bleached again and more colonies had died. In contrast, none of the previously unbleached colonies bleached at this time. In November 1999, just prior to the spawning period, there were large areas of coral on the reef slope that were noticeably pale. These pale colonies were sampled along with adjacent normally pigmented corals and the pale colonies were almost entirely devoid of eggs. During the 1998 bleaching event approximately 80% of the reef slope colonies were bleached, so it is likely that these pale colonies were previously bleached colonies. These data suggest that bleaching has adverse and long-lasting effects on coral reproduction and that previously bleached colonies may be more susceptible to future stress. Warner, ME, Fitt, WK and Schmidt, GW. (1999) Damage to photosystem I1 in symbiotic dinoflagellates: A determinant of coral bleaching. Proceedings of the National Academy of Sciences of the United States of America 96, 8007-6012. Coral bleaching has been defined as a general phenomenon, whereby reef corals turn visibly pale because of the loss of their symbiotic dinoflagellates and/or algal pigments during periods of exposure to elevated seawater temperatures. During the summer of 1997, seawater temperatures in the Florida Keys remained at or above 30°C for more than 6 weeks, and extensive coral bleaching was observed. Bleached colonies of the dominant Caribbean reefbuilding species, Montastrea faveolata and Montastrea franksi, were sampled over a depth gradient from 1to 17 m during this period of elevated temperature and contained lower densities of symbiotic dinoflagellates in deeper corals than seen in previous "nonbleaching" years. Fluorescence analysis by pulse-amplitude modulation fluorometry revealed severe

damage to photosystem I1 (PSII) in remaining symbionts within the corals, with greater damage indicated at deeper depths. Dinoflagellates with the greatest loss in PSII activity also showed a significant decline in the Dl reaction center protein of PSII, as measured by immunoblot analysis. Laboratory experiments on the temperature-sensitive species Montastrea annularis, as well as temperature-sensitive and temperature- tolerant cultured symbiotic dinoflagellates, confirmed the temperature-dependent loss of PSII activity and concomitant decrease in Dl reaction center protein seen in symbionts collected from corals naturally bleached on the reef. In addition, variation in PSII repair was detected, indicating that perturbation of PSII protein turnover rates during photoinhibition at elevated temperatures underlies the physiological collapse of symbionts in corals susceptible to heat-induced bleaching. Warner, ME, Chilcoat, GC, McFarland, FK and Fitt, WK. (2002) Seasonal fluctuations in the photosynthetic capacity of photosystem II in symbiotic dinoflagellates in the Caribbean reef-building coral Montastrea. Marine Biology 141, 31-38. The photosynthetic capacity of photosystem II (PS II) in symbiotic dinoflagellates (Symbiodinium sp.), as measured by analysis of chlorophyll fluorescence, was investigated in the primary Caribbean reef-building corals, Montastraea annularis and Montastraea faveolata, for 5 years and Montastraea franksi over 2 years in the Bahamas. Significant seasonal fluctuations in the quantum yield of charge separation (Fv/Fm) of PS II were found in all species at all depths, with the highest photosynthetic capacity consistently recorded between mid-winter and early spring and the lowest photosynthetic capacity occurring in the mid to late summer. Corals residing in shallow depths of 1–2 m showed the greatest fluctuations in Fv/Fm, whereas deeper corals (3– 4 and 14 m depths) had consistently higher values of Fv/Fm. Densities of symbiotic dinoflagellates and photosynthetic pigments followed a similar pattern. Fluctuations of photosynthetic capacity showed a strong correlation with seasonal patterns of water temperature and light. Such seasonal shifts in photosynthetic capacity are most likely due to several biochemical processes in the algae that lead to alterations of both photoprotection and photodamage. While symbiont density changed significantly on a seasonal basis, visual signs of coral bleaching were noted only in the fall of 1995 and the spring and summer of 1998. Comparisons of photosynthetic capacity and the decrease in the number of symbionts and their subsequent recovery indicated that symbiont populations in this study had the ability to recover quickly following bleaching events, as long as continued physical perturbation (e.g. thermal stress) did not shorten the recovery phase. Large-scale bleaching events are best viewed as the end points of seasonal physiological variation in which photosynthetic capacity and density of symbiotic dinoflagellates are reduced to a lower level than during ‘‘non-bleaching’’ years. West, JM and Salm, RV. (2003) Resistance and resilience to coral bleaching: Implications for coral reef conservation and management. Conservation Biology 17, 956-967. The massive scale of the 1997–1998 El Niño–associated coral bleaching event underscores the need for strategies to mitigate biodiversity losses resulting from temperature-induced coral mortality. As baseline sea surface temperatures continue to rise, climate change may represent the single greatest threat to coral reefs

worldwide. In response, one strategy might be to identify (1) specific reef areas where natural environmental conditions are likely to result in low or negligible temperature-related bleaching and mortality (i.e., areas of natural “resistance” to bleaching) and (2) reef areas where environmental conditions are likely to result in maximum recovery of reef communities after bleaching mortality has occurred (i.e., areas of natural community “resilience”). These “target areas,” where environmental conditions appear to boost resistance and resilience during and after large-scale bleaching events, could then be incorporated into strategic networks of marine protected areas designed to maximize conservation of global coral reef biodiversity. Based on evidence from the literature and systematically compiled observations from researchers in the field, this paper identifies likely environmental correlates of resistance and resilience to coral bleaching, including factors that reduce temperature stress, enhance water movement, decrease light stress, correlate with physiological tolerance, and provide physical or biological enhancement of recovery potential. As a tool for identifying reef areas that are likely to be most robust in the face of continuing climate change and for determining priority areas for reducing direct anthropogenic impacts, this information has important implications for coral reef conservation and management. Westmacott, S, Teleki, K, Wells, S and West, J. (2000) Management of bleached and severely damaged coral reefs. IUCN: Gland (Switzerland), 36 pp. This booklet was produced to provide guidance for managers, policy makers and all those who are concerned about the severe reef degradation caused by coral bleaching and a range of other impacts. Coral bleaching is caused by high sea surface temperatures and high levels of sunlight (UV), which affect the physiology of the coral and cause a whitening effect, or ‘bleaching’. This loss of colour is due to the loss of symbiotic algae (zooxanthellae) upon which the coral polyp depends for much of its food. Prolonged bleaching conditions (for over 10 weeks) can eventually lead to death of the coral polyp. Sustained high water temperatures (1–2°C above normal maximums) during 1998 caused the most geographically extensive bleaching event ever recorded. The Indian Ocean was one of the worst affected regions, with coral death as high as 90% over large areas of reef. The Pacific and Caribbean regions were also affected, but they did not experience the same level of coral mortality. Other human impacts continue to threaten the survival of coral reefs. Coastal development, poor land use practices, over exploitation of marine resources and destructive fishing methods — as well as waste disposal and pollution from ships — can all negatively affect the state of the reefs. Together, these impacts, especially when combined with increased coral bleaching, pose a serious threat to the survival of the world’s coral reefs. The Intergovernmental Panel on Climate Change (IPCC) has predicted an increase of 1–2°C in sea surface temperatures over the next 100 years, such that coral bleaching events will become a regular event in the next 30–50 years. Hence, the following types of management strategies will be crucial to safeguard coral reefs. 1. Marine Protected Areas (MPAs) will play a key role by helping to maintain sources of coral larvae to damaged areas. MPAs can also protect those areas where corals are struggling to recolonise damaged areas. Management actions in relation to MPAs, that will contribute to reef regeneration include:

• Identifying reef areas with least damage within MPAs and reviewing, and revising where necessary, zoning schemes and boundaries to ensure that healthy reefs are strictly protected. • Ensuring that existing MPAs are effectively managed. • Developing a more strategic approach to the establishment of MPA systems, including consideration of sources and sinks and inclusion of a wide geographic spread and variety of MPA types. 2. Reef fisheries may be negatively affected on reefs that have suffered major mortality and are losing their physical structure (and thus unable to support a diverse and abundant fish community). A precautionary approach can be taken by giving specific attention to the following: • Establishing no-fishing zones and limitations on fishing gear to protect breeding grounds and provide fish with a refuge. • Considering specific protection measures for species that can contribute to reef regeneration, such as algal grazers, or that might be affected by coral bleaching, such as coral-eating fishes. • Enforcing legislation prohibiting destructive fishing practices. • Monitoring the catch composition and size to evaluate the success of management strategies and implementing new strategies if necessary. • Developing alternative livelihoods for fishing communities as needed. • Limiting entry of new fishermen to a fishery through licensing schemes. • Regulating coral collection for the curio and aquarium trades. 3. Tourism in areas with bleached reefs can be maintained through the provision of other activities, both related and unrelated to the reef. Some management options include: • Maintaining healthy fish populations for divers and snorkellers through creative use of zoning to reduce pressure from overfishing and frequent tourist visitation. • Involving tourists in the bleaching issue by offering opportunities for participation in monitoring programmes. • Emphasising other attractions for tourists, both on land and in the water, besides coral reefs. • Reducing the impacts from tourism operations in general, such as direct damage to corals from divers and snorkellers or from boat anchors, and indirect damage from coastal activities that support the tourist industry. • Encouraging tourists to contribute financially to recovery and management efforts. • Conveying information to the public through outreach and education. 4. Integrated coastal management (ICM) will be crucial so that bleached reefs can be managed within the context of the land-use decisions being made in adjacent drainage basins. From the perspective of coral bleaching, particular aspects of ICM that need emphasising include: • Establishing MPA systems within an ICM framework. • Implementing measures to promote sustainable fisheries. • Implementing mechanisms to promote environmentally sound construction and other forms of land-use and coastal development. • Regulating land-based sources of pollution. • Managing shipping and other vessels to reduce damage to reefs from physical impacts or spills. • Protecting the coastline from erosion. 5. Reef restoration is a relatively new area of research. Research should be encouraged; however, costly rehabilitation programmes may be a risk rather than a

cure. Artificial rehabilitation should not be considered if human stressors continue to impact the reef. When considering restoration options, managers should consider the following questions: • What are the objectives of the restoration project? • What is the scale of the restoration project? • What will be the cost of the project, and is it affordable? • What is the success rate of the method being proposed, and which method will be most cost-effective at the site? • What will be the long term viability of the programme? • Is there scope for the local community and reef users to become involved? Monitoring will enable the managers and policy makers to track changes on the reef and assess the success of management programmes. Care must be taken to design a programme that fits within the personnel and financial capacity available. In many cases, there are existing programmes that can be adopted. Meanwhile, additional research is urgently needed so we can more fully answer key questions about the ecological and socio-economic impacts of coral bleaching. Managers can prepare for bleaching events and even aid reef recovery, but the global community needs to act now to tackle the issue of global climate change. Action at all levels from local communities and stakeholders to national governments and decision makers is required immediately to address not only the issues related to coral bleaching, but also the general state and plight of coral reefs everywhere. Wilkinson, C, Green, A, Almany, J and Dionne, S (eds.) (2003) Monitoring coral reef Marine Protected Areas. Australian Institute of Marine Science: Townsville (Australia), 68 pp. Marine Protected Areas (MPAs) are an important tool for marine conservation and management; monitoring plays a critical role in managing these MPAs. Monitoring provides the essential information required to make management decisions and determine if the decisions are working. Without monitoring, managers are essentially operating in the dark! This book was written in response to requests from many managers of MPAs from around the world who asked for advice on how to design and implement monitoring programs that can help them manage their MPAs more ffectively. The goals of this book are to: � Demonstrate how monitoring can play a major role in the effective management of MPAs; � Provide advice on which monitoring programs to use to facilitate effective management; and � Demonstrate how monitoring has played an important role in the effective management of MPAs using case studies from around the world. Coral reefs around the world are at risk from many threats including global warming causing coral bleaching, over-fishing or destructive fishing, pollution by sediments, nutrients and toxic chemicals, coral mining and shoreline development, and unregulated tourism. Monitoring the ecology of the reefs and the socioeconomics of the people is the only way to understand the extent, nature and causes of the damage, and to identify ways to address these threats. How can monitoring assist in the effective management of MPAs? Monitoring assists through the following tasks: 1. Resource Assessment and Mapping

2. Resource Status and Long-Term Trends 3. Status and Long-Term Trends of User Groups 4. Impacts of Large-Scale Disturbances 5. Impacts of Human Activities 6. Performance Evaluation and Adaptive Management 7. Education and Awareness Raising 8. Building Resilience into MPAs 9. Contributing to Regional and Global Networks This book will provide practical advice on how to design and implement ecological and socio-economic monitoring programs aimed at addressing these issues. Many useful references are included at the back along with Internet sites. We have used case studies from around the world to illustrate how others have used monitoring to assist them in managing MPAs. There are many useful lessons from these case studies and all contain recommendations for other MPA managers. The book provides information on many of the organisations involved in coral reef monitoring and management, along with the recommendations on coral reef monitoring and information processing from the recent ITMEMS2 (International Tropical Marine Ecosystems Management Symposium, 2003) meeting, which featured MPA managers from all over the world. This is Version 1 of the book being released at the World Parks Congress in Durban South Africa, September 2003. Our intention is to keep it alive and continually update it. This copy will be lodged on the www.reefbase.org, www.gcrmn.org and www.aims.gov.au websites where we want to continually update it for use by MPA managers to improve their management and conservation of coral reefs. Wilkinson, C (ed.) (2004) Status of coral reefs of the world: 2004. Australian Institute of Marine Science: Townsville (Australia), 557 pp. The assessments of more than 240 contributors from 98 countries in this Status of Coral Reefs of the World: 2004 report that: CURRENT STATUS OF CORAL REEFS - Estimates in this report are that 20% of the world’s coral reefs have been effectively destroyed and show no immediate prospects of recovery; - Approximately 40% of the 16% of the world’s reefs that were seriously damaged in 1998 are either recovering well or have recovered; - The report predicts that 24% of the world’s reefs are under imminent risk of collapse through human pressures; and a further 26% are under a longer term threat of collapse; - Coral reefs around the world continue to decline from increasing human pressures; poor land management practices are releasing more sediment, nutrients and other pollutants that stress reefs; - Over-fishing and particularly fishing with destructive methods are: threatening the normal functioning of coral reef ecosystems; reducing populations of key reef organisms; lowering coral reef productivity; and, along with pollution, shift the advantage towards macro-algae by removing grazing pressure. These algae smother and out-compete corals; - Pressures on reefs from coral predators such as the crown-of-thorns starfish (COTS) and coral disease have not increased recently (sometimes because corals have declined); but severe problems remain on some reefs. There is evidence that these are exacerbated by human pressures, either by removing the predators of

COTS and/or increasing water temperatures that stress corals, making them more susceptible to coral diseases; - Analyses of coral reefs in the wider Caribbean region confirm major reef declines and they do not resemble the reefs of 30 years ago. Coral cover on many Caribbean reefs has declined by up to 80%; however there are some encouraging signs of recovery; - There are few encouraging signs for reefs in the high biodiversity areas of Southeast Asia and the Indian Ocean, where human pressures continue to increase on coral reef; whereas reefs in the Pacific and around Australia remain quite healthy. GLOBAL THREATS TO CORAL REEFS - Many coral reefs continue to recover after the 1998 El Niño/La Niña global coral bleaching event, with stronger recovery in well-managed and remote reefs; however, the recovery is not uniform and many reefs virtually destroyed in 1998 show minimal signs of recovery. This recovery could be reversed if the predicted increases in ocean temperatures occur as a result of increasing global climate change; - There has been no recurrence of the major global-scale climate change pressures of 1998; although there have been some more localised bleaching events in 2000 and 2003 causing damage to reefs; - The coral bleaching in 1998 was a 1 in a 1000-year event in many regions with no past history of such damage in official government records or in the memories of traditional cultures of the affected coral reef countries. Also very old corals around 1000 years old died during 1998. Increasing sea surface temperatures and CO2 concentrations provide clear evidence of global climate change in the tropics, and current predictions are that the extreme events of 1998 will become more common in the next 50 years, i.e. massive global bleaching mortality will not be a 1/1000 year event in the future, but a regular event; - Coral disease and major coral predators like the crown-of-thorns starfish continue to threaten reefs and evidence points to human disturbance as a contributing and catalytic factor behind these increases. CORAL REEF MANAGEMENT, AWARENESS RAISING AND POLITICAL WILL - There was a major advance in the protection of the Great Barrier Reef with increases in the amount of no-take areas from 5% to 33%, following a careful analysis using the best available science and extensive consultation with major stakeholders; - The World Summit on Sustainable Development in 2002 called for the establishment of networks of larger marine protected areas (MPAs) and a major international effort to reduce losses in biodiversity, including the biodiversity on tropical and cold-water coral reefs; - Many coral reef countries lack the resources of trained personnel, equipment and finances to effectively conserve coral reefs, establish MPAs and enforce regulations; - This lack of resources is often exacerbated by a poor awareness of the problems facing coral reefs and their significance in local economies, and inadequate political will to tackle difficult environmental problems; - Major international NGOs are combining their expertise and resources to establish networks of MPAs and improve management capacity. A major focus is on the high biodiversity region of Southeast Asia and the Western Pacific; - Some of these NGOs have developed rapid assessment methods to select sites for urgent protection and also designed tools to assist resource managers protect reefs from global change stresses;

- International interest and political will for the protection of coral reefs is improving and the International Coral Reef Initiative is expanding to further catalyse improved management of coral reefs and raise the profile of all coral reefs within global forums; - Cold water coral reefs are now being recognised as valuable resources that warrant protection from the massive degradation being caused to them by deep water trawling; and - Millions of people around the world were made aware of coral reef conservation via the animated film on Nemo on the Great Barrier Reef; the film carried many coral reef conservation messages, but there were unfortunate consequences with an increase the trade of aquarium species and the release of some species in the wrong regions. Williams, ID and Polunin, NVC. (2001) Large-scale associations between macroalgal cover and grazer biomass on mid-depth reefs in the Caribbean. Coral Reefs 19, 358-366. Since the 1970s, macroalgae have become considerably more abundant on many Caribbean reefs and overfishing of grazing fishes has been implicated as a contributory factor. We explored relationships between algal cover and grazers (biomass of herbivorous fishes and abundance of the sea-urchin Diadema antillarum) on mid-depth reefs (12-15 m) in 19 areas at seven locations in Jamaica Barbados, Belize, Grand Cayman and Cuba between April 1997 and April 1998. Diadema antillarum density was never >0.01 m-2, while herbivorous fish biomass (acanthurids and scarids ≥12 cm total length) varied from 2-5 g m-2 in Jamaica to 17.1 g m-2 in Barbados, and was strongly correlated, negatively with macroalgal cover and positively with cropped' substratum (sum of bare', turf and crustose-coralline substrata) cover. However, overfishing of herbivorous fishes alone cannot explain the widespread abundance of macroalgae, as even on lightly fished reefs, macroalgal cover was mostly > 20%. Herbivorous fish populations on those reefs were apparently only able to maintain approximately 40-60% of reef substratum in cropped states, but due to low space-occupation by coral and other invertebrates, 70-90% of substratum was available to algae. The abundance of macroalgae on lightly fished reefs may therefore be a symptom of low coral cover in combination with the continuing absence of Diadema antillarum. Willis, BL, van Oppen, MJH, Miller, DJ, Vollmer, SV and Ayre, DJ. (2006) The Role of Hybridization in the Evolution of Reef Corals. Annual Review of Ecology, Evolution and Systematics 37, 489-517. The importance of hybridization in the evolution of plant species is widely accepted, but its contributions to animal species evolution remain less recognized. Here we review evidence that hybridization has contributed to the evolution of reef corals, a group underpinning the coral reef ecosystem. Increasingly threatened by human and climate-related impacts, there is need to understand the evolutionary processes that have given rise to their diversity and contribute to their resilience. Reticulate evolutionary pathways among the ecologically prominent, mass-spawning genus Acropora suggest that hybridization, although rare on ecological timescales, has been instrumental in their diversification on evolutionary timescales. Evidence that coral hybrids colonize marginal habitats distinct from those of parental species’ and that hybridization may be more frequent at peripheral boundaries of species’ ranges supports a role for hybridization in range expansion and adaptation to changing

environments. We conclude that outcomes of hybridization are significant for the future resilience of reef corals and warrant inclusion in conservation strategies. Wilson, J and Harrison, P. (2005) Post-settlement mortality and growth of newly settled reef corals in a subtropical environment. Coral Reefs 24, 418–421. There are few studies of post-settlement mortality of newly settled corals, particularly on subtropical reefs. This study examined the mortality and growth of spat of three broadcast spawning coral species at the Solitary Islands, eastern Australia (30°S). Mortality of spat was high, with only 0.2–2.8% surviving their first year of life. Growth was slow, with coral spat achieving a maximum diameter of 2 mm after 8 months. High post-settlement mortality and slow growth rates are likely to contribute to low rates of recruitment of broadcast spawning species at the Solitary Islands. Wilson, SK, Graham, NAJ, Pratchett, MS, Jones, GP and Polunin, NVC. (2006) Multiple disturbances and the global degradation of coral reefs: are reef fishes at risk or resilient? Global Change Biology 12, 2220–2234. Increased frequency of disturbances and anthropogenic activities are predicted to have a devastating impact on coral reefs that will ultimately change the composition of reef associated fish communities. We reviewed and analysed studies that document the effects of disturbance-mediated coral loss on coral reef fishes. Meta-analysis of 17 independent studies revealed that 62% of fish species declined in abundance within 3 years of disturbances that resulted in 410% decline in coral cover. Abundances of species reliant on live coral for food and shelter consistently declined during this time frame, while abundance of some species that feed on invertebrates, algae and/or detritus increased. The response of species, particularly those expected to benefit from the immediate loss of coral, is, however, variable and is attributed to erratic replenishment of stocks, ecological versatility of species and sublethal responses, such as changes in growth, body condition and feeding rates. The diversity of fish communities was found to be negatively and linearly correlated to disturbance-mediated coral loss. Coral loss 420% typically resulted in a decline in species richness of fish communities, although diversity may initially increase following small declines in coral cover from high coverage. Disturbances that result in an immediate loss of habitat complexity (e.g. severe tropical storms), have a greater impact on fishes from all trophic levels, compared with disturbances that kill corals, but leave the reef framework intact (e.g. coral bleaching and outbreaks of Acanthaster planci). This is most evident among small bodied species and suggests the long-term consequences of coral loss through coral bleaching and crown-of-thorn starfish outbreaks may be much more substantial than the short-term effects currently documented. Winters, G, Loya, Y and Beer, S. (2006) In situ measured seasonal variations in Fv/Fm of two common Red Sea corals. Coral Reefs 25: 593–598. Pulse amplitude modulated (PAM) fluorometry has been suggested as a tool for estimating environmental stresses on corals. However, information regarding natural changes in maximal quantum yields (Fv/Fm) of corals during “normal” (i.e. non-bleaching) years has been limited. In this study, seasonal variations in Fv/Fm for Stylophora pistillata and Favia favus, measured in situ, correlated with seasonal

changes in solar irradiance but not in sea temperature. Interactions between sea temperature and irradiance were further studied by growing these corals and Pocillopora damicornis under controlled conditions. Exposure to high light with normal or high temperatures resulted in lower Fv/Fm values than exposure to low light at both temperatures. Thus, high irradiances may cause decreased Fv/Fm values in corals at least as much as, if not more than, high temperatures. Such seasonal variations should be taken into account when using PAM fluorometry as a diagnostic tool for predicting coral bleaching. Wooldridge, S and Done, T. (2004) Learning to predict large-scale coral bleaching from past events: A Bayesian approach using remotely sensed data, in-situ data, and environmental proxies. Coral Reefs 23, 96-108. Ocean warming and coral bleaching are patchy phenomena over a wide range of scales. This paper is part of a larger study that aims to understand the relationship between heat stress and ecological impact caused by the 2002-bleaching event in the Great Barrier Reef (GBR). We used a Bayesian belief network (BBN) as a framework to refine our prior beliefs and investigate dependencies among a series of proxies that attempt to characterize potential drivers and responses: the remotely sensed environmental stress (sea surface temperature — SST); the geographic setting; and topographic and ecological attributes of reef sites for which we had field data on bleaching impact. Sensitivity analyses helped us to refine and update our beliefs in a manner that improved our capacity to hindcast areas of high and low bleaching impact. Our best predictive capacity came by combining proxies for a site’s heat stress in 2002 (remotely sensed), acclimatization temperatures (remote sensed), the ease with which it could be cooled by tidal mixing (modeled), and type of coral community present at a sample of survey sites (field data). The potential for the outlined methodology to deliver a transparent decision support tool to aid in the process of identifying a series of locations whose inclusion in a network of protected areas would help to spread the risk of bleaching is discussed. WWF (2003) Securing Australia’s Great Barrier Reef. WWF: Sydney (Australia), 21 pp. Throughout Australia’s history, the Great Barrier Reef has been a source of wonder, awe and inspiration. Its unparalleled beauty now inspires people the world over, and it continues to provide wealth through tourism and fishing. Until recently, the Great Barrier Reef was regarded as a well-protected, pristine wonderland. A place of delicate corals, abundant fish life and a haven for other marine life. As scientists have understood more about the Reef’s complexities, a different picture has emerged of the Reef’s brilliance being tarnished by overfishing, land-based pollution and coral bleaching exacerbated by increased sea temperatures due to global warming. The Reef is unlikely to die in the coming decades. However, it is likely to change significantly from the system that we now know. The good news is that due to its remoteness from the coastline, its size and Australia’s small and generally affluent population, the Great Barrier Reef hasn’t suffered from the extreme human impacts that have caused catastrophic damage to many other coral reefs around the world. We can protect the Reef. Australians have a far better chance of giving our coral reefs to future generations in a healthy state, than anywhere else in the world. It’s not too late. In early 2003, 94% of Australians

(including more than 90% of coastal Queenslanders) said they wanted greater protection for the Great Barrier Reef.2 Currently, only 4.6% of the entire Marine Park is highly protected in green zones (places where commercial and recreational fishing are prohibited). Aside from its unparalleled biological riches, the Great Barrier Reef is also the source of enormous economic wealth. Tens of thousands of Australian families depend on the Reef to maintain their way of life. Tourism is by far the biggest Reef-based industry. The Great Barrier Reef attracts in excess of 1.6 million visitors annually and it is consistently rated as one of the top three attractions for visitors coming to Australia.3 Tourism contributes $4.2 billion to the gross value of production in the Great Barrier Reef catchment and is by far the largest employer with 47,660 employees. “The health and future of Reef-based tourism is dependent on the long-term health and future of the Great Barrier Reef.” By comparison, recreational fishing contributes around $187 million and commercial fishing is estimated to add another $118 million to the regional economies along the Reef coastline. The Australian and Queensland governments have recognised the biological and economic importance of the Great Barrier Reef, and the threats it faces. They are putting in place the following initiatives: • The Representative Areas Program: an Australian government program to protect the biodiversity of the Great Barrier Reef World Heritage area by rezoning the Marine Park to establish a comprehensive network of green zones; and • The Reef Water Quality Protection Plan: a joint Australian and Queensland government plan to halt and reverse the decline in the quality of water flowing into the Reef caused by poor land management practices in the catchments adjacent to the Great Barrier Reef World Heritage area. “The Representative Areas Program and the Reef Water Quality Protection Plan are both critical to maintaining and restoring the health of the Reef. They therefore underpin the future health of the tourism industry in Queensland.” State fisheries management plans are also slowly being developed to ensure that the various fisheries in the Marine Park are ecologically sustainable. This document presents WWF Australia’s proposal for a world class network of green zones throughout the Great Barrier Reef Marine Park. We describe why greater protection is beneficial and where additional green zones should be placed. Our full submission to the Representative Areas Program can be found at www.gbr.wwf.org.au. Yakovleva, I and Hidaka, M. (2004) Diel fluctuations of mycosporine-like amino acids in shallow-water scleractinian corals. Marine Biology 145, 863-873. Reports of bathymetric decrease in the total mycosporine-like amino acid (MAA) concentration of benthic invertebrates suggest that light gradients may be important determinants of MAA content. With the pronounced diel light changes, distinct temporal variations in MAA concentrations might also be expected. We examined the changes in the abundance of MAA in three shallow-water scleractinian corals, Pavona divaricata, Galaxea fascicularis and Montipora digitata from Okinawa, Japan, in relation to daily cycles in solar radiation and tested whether the species have different capabilities for protection against UVR depending on their MAA composition. The results show that symbiotic algae freshly isolated from the investigated coral species do not contain MAAs and that distribution of these compounds resided only within the animal tissue. Total MAA content in the tissue of P. divaricata, G. fascicularis and M. digitata rose rapidly at midday and significantly dropped at night.

The observed variations were by a factor of two and, thus, very dramatic. For all the investigated coral species, total MAA concentrations were significantly correlated with the diurnal cycle in solar radiation, during both winter and summer seasons. Seawater temperature was significantly correlated with MAA levels only in the June experiment, but represented no more than 20% of the MAA variation in all three coral species, whereas solar radiation explained 60–70% of the MAA fluctuations. This suggests that MAAs are an integral component of the hard coral’s biochemical defense system against high solar irradiance stress. The diurnal increase in total MAA concentrations was due to an increase in the concentration of imino-MAA species of up to 2–2.5-fold of their pre-dawn values. In contrast, the oxocarbonyl-MAA mycosporine-glycine (Myc-Gly) showed the lowest (Tukey–Kramer HSD test: P<0.05) values at midday, compared to afternoon and night hours. Analysis of diel changes in chlorophyll fluorescence and chlorophyll a content of the investigated species revealed that P. divaricata and G. fascicularis were less sensitive to the high levels of ambient irradiance compared to M. digitata. In P. divaricata and G. fascicularis, Myc-Gly, an MAA with an antioxidant function, is the most abundant MAA, contributing about 70% to the total MAA pool, whereas the major MAA factions in M. digitata were represented by oxidatively robust imino-MAAs. We speculate that MAAs furnish scleractinian corals with protection from biologically damaging ultraviolet radiation through both the direct sunscreening activity of imino-MAAs and the antioxidant properties of oxocarbonyl-MAAs and suggest that the predominance, in the host tissue, of MAAspecies with an antioxidant ability may render corals more tolerant to high photosynthetically active and ultraviolet radiation. Yakovleva, I, Bhagooli, R, Takemura, A and Hidaka, M. (2004) Differential susceptibility to oxidative stress of two scleractinian corals: Antioxidant functioning of mycosporine-glycine. Comparative Biochemistry and Physiology. Part B Biochemistry and Molecular Biology 139, 721-730. This study examined the importance of mycosporine-glycine (Myc-Gly) as a functional antioxidant in the thermal-stress susceptibility of two scleractinian corals, Platygyra ryukyuensis and Stylophora pistillata. Photochemical efficiency of PSII (F(v)/F(m)), activity of antioxidant enzymes, superoxide dismutase (SOD) and catalase (CAT), and composition and abundance of mycosporine-like amino acids (MAAs) in the coral tissue and in symbiotic zooxanthellae were analyzed during 12-h exposure to high temperature (33 degrees C). After 6- and 12-h exposures at 33 degrees C, S. pistillata showed a significantly more pronounced decline in F(v)/F(m) compared to P. ryukyuensis. A 6-h exposure at 33 degrees C induced a significant increase in the activities of SOD and CAT in both host and zooxanthellae components of S. pistillata while in P. ryukyuensis a significant increase was observed only in the CAT activity of zooxanthellae. After 12-h exposure, the SOD activity of P. ryukyuensis was unaffected in the coral tissue but slightly increased in zooxanthellae, whereas the CAT activity in the coral tissue showed a 2.5-fold increase. The total activity of antioxidant enzymes was significantly higher in S. pistillata than in P. ryukyuensis, suggesting that P. ryukyuensis is less sensitive to oxidative stress than S. pistillata. This differential susceptibility of the corals is consistent with a 20-fold higher initial concentration of Myc-Gly in P. ryukyuensis compared to S. pistillata. In the coral tissue and zooxanthellae of both species investigated, the first 6 h of exposure to thermal stress induced a pronounced reduction in the abundance of Myc-Gly but not in other MAAs. When exposure was

prolonged to 12 h, the Myc-Gly pool continued to decrease in P. ryukyuensis and was completely depleted in S. pistillata. The delay in the onset of oxidative stress in P. ryukyuensis and the dramatic increase in the activities of the antioxidant enzymes in S. pistillata, which contains low concentrations of Myc-Gly suggest that Myc-Gly provides rapid protection against oxidative stress before the antioxidant enzymes are induced. These findings strongly suggest that Myc-Gly is functioning as a biological antioxidant in the coral tissue and zooxanthellae and demonstrate its importance in the survival of reef-building corals under thermal stress. Yamano, H and Tamura, M. (2004) Detection limits of coral reef bleaching by satellite remote sensing: Simulation and data analysis. Remote Sensing of Environment 90, 86-103. Monitoring of coral reef bleaching has hitherto been based on regional-scale, in situ data. Larger-scale trends, however, must be determined using satellite-based observations. Using both a radiative transfer simulation and an analysis of multitemporal Landsat TM images, the ability of satellite remote sensing to detect and monitor coral reef bleaching is examined. The radiative transfer simulation indicates that the blue and green bands of Landsat TM can detect bleaching if at least 23% of the coral surface in a pixel has been bleached, assuming a Landsat TM pixel with a resolution of 30x30 m on shallow (less than 3 m deep) reef flats at Ishigaki Island, Japan. Assuming an area with an initial coral coverage of 100% and in which all corals became completely bleached, the bleaching could be detected at a depth of up to 17 m. The difference in reflectance of shallow sand and corals is compared by examining multitemporal Landsat TM images at Ishigaki Island, after normalizing for variations in atmospheric conditions, incident light, water depth, and the sensor’s reaction to the radiance received. After the normalization, a severe bleaching event when 25–55% of coral coverage was bleached was detected, but a slight bleaching event when 15% of coral coverage was bleached was not detected. The simulation and data analysis agreed well with each other, and identified reliable limits for satellite remote sensing for detecting coral reef bleaching. Sensitivity analysis on solar zenith angle, aerosol (visibility) and water quality (Chl a concentration) quantified the effect of these factors on bleaching detection, and thus served as general guidelines for detecting coral reef bleaching. Spatial misregistration resulted in a high degree of uncertainty in the detection of changes at the edges of coral patches mainly because of the low (f30 m) spatial resolution of Landsat TM, indicating that detection of coral reef bleaching by Landsat TM is limited to extremely severe cases on a large homogeneous coral patch and shallow water depths. Satellite remote sensing of coral reef bleaching should be encouraged, however, because the development and deployment of advanced satellite sensors with high spatial resolution continue to progress. Yap, HT, Alvarez, RM, Custodio III, HM and Dizon, RM. (1998) Physiological and ecological aspects of coral transplantation. Journal of Experimental Marine Biology and Ecology 229, 69-84. The growth and mortality of transplants of two species of scleractinian corals, Porites cylindrical Dana and P. rus Forskal (1775), were monitored over 16 months in a reef in the northwestern Philippines. Transplants were in two sizes (nubbin, ~8 cm in length; and fist-sized, ~8 cm in diameter) and deployed at two depths (1 and 10 m).

Specimens at the shallow depth had more rapid growth than the deeper ones. Light had significant effects on coral growth while temperature, salinity, water motion and sedimentation did not. Smaller corals consistently registered greater percentage increases in size as compared to the larger ones. Small transplants of P. cylindrica showed higher percentage growth rates than those of P. rus. Contrary to expectations, there were no differences in transplant mortality due to size. Over the course of the experiment, mortality was generally confined to the shallow depth. It was brought about either by algal competition or by strong water movement. Yap, HT. (2000) The case for restoration of tropical coastal ecosystems. Ocean and Coastal Management 43, 841-851. At no time have humans so altered their natural environment than the present. Marine ecosystems have not been spared, and the degradation of coastal habitats has reached severe proportions in many parts of the world. The mere setting aside of areas for protection may not be enough to ensure adequate production and provision of services for a growing global human population. Hence, the active restoration of habitats, in addition to protection and preservation, is probably the more desirable conservation strategy. Accumulated experience over several decades has demonstrated that the rehabilitation or even restoration of damaged coastal ecosystems is feasible. However, the degree of difficulty and expense involved vary, with coral reefs being the most complicated habitats to restore, followed by seagrass beds and then mangrove forests. In ecosystem restoration, a comprehensive strategy based on sound biological and ecological principles, and proven techniques must be developed. A concrete, achievable goal must be articulated. Because of the dynamic nature of ecosystems, and the inability to accurately predict pathways of succession after a community is established through artificial means, subsequent modifications to a project must proceed within a flexible framework of adaptive management. Finally, for restoration efforts to be successful, local communities must participate actively in cooperation with local governments in accordance with the principle of co-management. Yap, HT. (2004) Differential survival of coral transplants on various substrates under elevated water temperatures. Marine Pollution Bulletin 49, 306-312. Closely related scleractinian coral species that exhibited similar survival patterns under relatively normal field conditions responded very differently to the occurrence of an environmental disturbance. The two species studied were Porites cylindrica and Porites rus which occur in the same reef zones in shallow reef flats. Transplants of both species were evenly distributed and attached to three different types of substrate: live coral colonies of P. cylindrica, dead coral colonies (also of P. cylindrica), and epoxy coated metal grids that were raised above the sandy substrate. With the onset of above-normal water temperatures due to the El Niño episode of 1998, P. cylindrica transplants immediately showed signs of bleaching stress and tissue necrosis, followed by algal overgrowth and mortality soon afterwards. In contrast, transplants of P. rus bleached more slowly and suffered less mortality, with a few actually showing signs of recovery at the end of the experimental period which covered a total of 14 weeks. These differences in responses could be attributed to properties of the symbiotic zooxanthellae, of the host coral tissue itself, or both. Overall, survival was good on the metal grids (average of 35%), and on the live coral

(average of 22%). It was poor on the dead coral (average of 6%). The metal grids as well as live coral tissue apparently provided a favorable substrate for the attached coral fragments, even for those of a different species. Under the conditions of this particular study, attachment of live coral fragments on already dead colonies for the purpose of increasing live coral cover on the reef did not yield favorable results. This is an area that requires further investigation. Yap, HT. (2007) Coral reef resilience. Marine Pollution Bulletin 54, 1075–1076. Coral reef ecosystems are vulnerable to catastrophic collapse, but can also be resilient or robust. This proposition was reinforced, first-hand, during a recent inspection of a reef slope in Bolinao, Pangasinan in the northwestern Philippines, which traditionally has been a favorite diving spot. During the 1998 severe bleaching event, this entire reef slope, covering hundreds of square meters, turned stark white. It appeared no different from a snow-covered mountain slope! The following year, almost all visible coral was dead and/or overgrown by filamentous algae. It was a depressing sight, indeed. However, over the following years, coral recruitment, vigorous in some places, was observed to be taking place. Now, many parts of that reef slope are again thriving with a variety of hard coral species. Except for the occasional, tell-tale dead colonies that are still around, it would be virtually impossible to tell that catastrophic mortality had taken place almost a decade previously. We have not established where the coral recruits might be coming from, but obviously there are enough sources in the vicinity, connected to the reef by water currents. Some of the living coral cover also probably grew back from remnants on colonies that did not perish entirely from the bleaching. Yentsch, CS, Yentsch, CM, Cullen, JJ, Lapointe, B, Phinney, DA and Yentsch, SW. (2002) Sunlight and water transparency: Cornerstones in coral research. Journal of Experimental Marine Biology and Ecology 268, 171-183. Reef-building corals throughout the world are considered endangered. The evidence is a decline in coral health and reduced coral cover. Competing hypotheses for the cause of coral loss include removal of grazers, nutrient enrichment, disease, coral bleaching, increase in temperature, and excess light/ultraviolet exposure. We suggest that light limitation as a second order effect of anthropogenic activity (e.g. sediment resuspension and nutrient enrichment) is a valid and tractable hypothesis. This experimental field and laboratory study demonstrates that corals of the Florida reefs are functioning close to the compensation point where respiration (of coral polyp plus zooxanthellae) consumes the products of photosynthesis of the zooxanthellae, with little if any remaining for growth. We extend this work into an optical nomograph that is useful for predicting coral loss and recovery. The nomograph is designed to elucidate compensation depth for waters of various transparencies.