C2.2.1 Are coral reefs endangered by climate change? (Chapter 4, Box 4.4)
Reefs are habitat for about a quarter of all marine species and are the most diverse among marine ecosystems (Roberts et al., 2002; Buddemeier et al., 2004). They underpin local shore protection, fisheries, tourism (see Chapter 6; Hoegh-Guldberg et al., 2000; Cesar et al., 2003; Willig et al., 2003; Hoegh-Guldberg, 2004,2005) and, although supplying only about 2-5% of the global fisheries harvest, comprise a critical subsistence protein and income source in the developing world (Whittingham et al., 2003; Pauly et al., 2005; Sadovy, 2005).
Corals are affected by warming of surface waters (see C2.1.2; Reynaud et al., 2003; McNeil et al., 2004; McWilliams et al., 2005) leading to bleaching (loss of algal symbionts; see C2.1.2). Many studies incontrovertibly link coral bleaching to warmer sea surface temperature (e.g., McWilliams et al., 2005), and mass bleaching and coral mortality often results beyond key temperature thresholds (see C2.1.2). Annual or bi-annual exceedance of
1 A clade of algae is a group of closely related, but nevertheless different, types.
Figure C2.2. Alternative hypotheses concerning the threshold SSTat which coral bleaching occurs: (a) invariant threshold for coral bleaching (red line) which occurs when SST exceeds usual seasonal maximum threshold (by ~1°C) and mortality (dashed red line, threshold of2°C), with local variation due to different species or water depth; (b) elevated threshold for bleaching (green line) and mortality (dashed green line) where corals adapt or acclimatise to increased SST (based on Hughes et al., 2003).
bleaching thresholds is projected at the majority of reefs worldwide by 2030 to 2050 (Hoegh-Guldberg, 1999; Sheppard, 2003; Donner et al., 2005). After bleaching, algae quickly colonise dead corals, possibly inhibiting later coral recruitment (e.g., McClanahan et al., 2001; Szmant, 2001; Gardner et al., 2003; Jompa and McCook, 2003). Modelling predicts a phase switch to algal dominance on the Great Barrier Reef and Caribbean reefs in 2030 to 2050 (Wooldridge et al., 2005).
Coral reefs will also be affected by rising atmospheric CO2 concentrations (Orr et al., 2005; Raven et al., 2005; Denman et al., 2007, Box 7.3) resulting in declining calcification. Experiments at expected aragonite concentrations demonstrated a reduction in coral calcification (Marubini et al., 2001; Langdon et al., 2003; Hallock, 2005), coral skeleton weakening (Marubini et al., 2003) and strong temperature dependence (Reynaud et al., 2003). Oceanic pH projections decrease at a greater rate and to a lower level than experienced over the past 20 million years (Caldeira and Wickett, 2003; Raven et al., 2005; Turley et al., 2006). Doubling CO2 will reduce calcification in aragonitic corals by 20%-60% (Kleypas et al., 1999; Kleypas and Langdon, 2002; Reynaud et al., 2003; Raven et al., 2005). By 2070 many reefs could reach critical aragonite saturation states (Feely et al., 2004; Orr et al., 2005), resulting in reduced coral cover and greater erosion of reef frameworks (Kleypas et al., 2001; Guinotte et al., 2003).
Adaptation potential (Hughes et al., 2003) by reef organisms requires further experimental and applied study (Coles and Brown, 2003; Hughes et al., 2003). Natural adaptive shifts to symbionts with +2°C resistance may delay the demise of some reefs until roughly 2100 (Sheppard, 2003), rather than mid-century (Hoegh-Guldberg, 2005) although this may vary widely across the globe (Donner et al., 2005). Estimates of warm-water coral cover reduction in the last 20-25 years are 30% or higher (Wilkinson, 2004; Hoegh-Guldberg, 2005) due largely to increasing higher SST frequency (Hoegh-Guldberg, 1999). In some regions, such as the Caribbean, coral losses have been estimated at 80% (Gardner et al., 2003). Coral migration to higher latitudes with more optimal SST is unlikely, due both to latitudinally decreasing aragonite concentrations and projected atmospheric CO2 increases (Kleypas et al., 2001; Guinotte et al., 2003; Orr et al., 2005; Raven et al., 2005). Coral migration is also limited by lack of available substrate (see C2.2.2). Elevated SST and decreasing aragonite have a complex synergy (Harvell et al., 2002; Reynaud et al., 2003; McNeil et al., 2004; Kleypas et al., 2005) but could produce major coral reef changes (Guinotte et al., 2003; Hoegh-Guldberg, 2005). Corals could become rare on tropical and sub-tropical reefs by 2050 due to the combined effects of increasing CO2 and increasing frequency of bleaching events (at 2-3 x CO2) (Kleypas and Langdon, 2002; Hoegh-Guldberg, 2005; Raven et al., 2005). Other climate change factors (such as sea-level rise, storm impact and aerosols) and non-climate factors (such as over-fishing, invasion of non-native species, pollution, nutrient and sediment load (although this could also be related to climate change through changes to precipitation and river flow; see C2.1.2 and C2.2.3; Chapter 16)) add multiple impacts on coral reefs (see C2.3.1), increasing their vulnerability and reducing resilience to climate change (Koop et al., 2001; Kleypas and Langdon, 2002; Cole, 2003; Buddemeier et al., 2004; Hallock, 2005).
C2.2.2 Impacts on coral reefs (Chapter 6, Section 126.96.36.199)
Reef-building corals are under stress on many coastlines (see C2.1.1). Reefs have deteriorated as a result of a combination of anthropogenic impacts such as over-fishing and pollution from adjacent land masses (Pandolfi et al., 2003; Graham et al., 2006), together with an increased frequency and severity of bleaching associated with climate change (see C2.1.2). The relative significance of these stresses varies from site to site. Coral mortality on Caribbean reefs is generally related to recent disease outbreaks, variations in herbivory,2 and hurricanes (Gardner et al., 2003; McWilliams et al., 2005), whereas Pacific reefs have been particularly impacted by episodes of coral
2 Herbivory: the consumption of plants by animals. 852
bleaching caused by thermal stress anomalies, especially during recent El Niño events (Hughes et al., 2003), as well as non-climate stresses.
Mass coral-bleaching events are clearly correlated with rises of SST of short duration above summer maxima (Douglas, 2003; Lesser, 2004; McWilliams et al., 2005). Particularly extensive bleaching was recorded across the Indian Ocean region associated with extreme El Niño conditions in 1998 (see C2.1.2 and C2.2.3). Many reefs appear to have experienced similar SST conditions earlier in the 20th century and it is unclear how extensive bleaching was before widespread reporting post-1980 (Barton and Casey, 2005). There is limited ecological and genetic evidence for adaptation of corals to warmer conditions (see C2.1.2 and C2.2.1). It is very likely that projected future increases in SST of about 1 to 3°C (Section 6.3.2) will result in more frequent bleaching events and widespread mortality if there is no thermal adaptation or acclimatisation by corals and their symbionts (Sheppard, 2003; Hoegh-Guldberg, 2004). The ability of coral reef ecosystems to withstand the impacts of climate change will depend on the extent of degradation from other anthropogenic pressures and the frequency of future bleaching events (Donner et al., 2005).
In addition to coral bleaching, there are other threats to reefs associated with climate change (Kleypas and Langdon, 2002). Increased concentrations of CO2 in seawater will lead to ocean acidification (Section 6.3.2), affecting aragonite saturation state (Meehl et al., 2007) and reducing calcification rates of calcifying organisms such as corals (LeClerq et al., 2002; Guinotte et al., 2003; see C2.2.1). Cores from long-lived massive corals indicate past minor variations in calcification (Lough and Barnes, 2000), but disintegration of degraded reefs following bleaching or reduced calcification may result in increased wave energy across reef flats with potential for shoreline erosion (Sheppard et al., 2005). Relative sea-level rise appears unlikely to threaten reefs in the next few decades; coral reefs have been shown to keep pace with rapid postglacial sea-level rise when not subjected to environmental or anthropogenic stresses (Hallock, 2005). A slight rise in sea level is likely to result in the submergence of some Indo-Pacific reef flats and recolonisation by corals, as these intertidal surfaces, presently emerged at low tide, become suitable for coral growth (Buddemeier et al., 2004).
Many reefs are affected by tropical cyclones (hurricanes, typhoons); impacts range from minor breakage of fragile corals to destruction of the majority of corals on a reef and deposition of debris as coarse storm ridges. Such storms represent major perturbations, affecting species composition and abundance, from which reef ecosystems require time to recover. The sequence of ridges deposited on the reef top can provide a record of past storm history (Hayne and Chappell, 2001); for the northern Great Barrier Reef no change in frequency of extremely large cyclones has been detected over the past 5,000 years (Nott and Hayne, 2001). An intensification of tropical storms (Section 6.3.2) could have devastating consequences on the reefs themselves, as well as for the inhabitants of many low-lying islands (Sections 6.4.2 and 188.8.131.52). There is limited evidence that global warming may result in an increase of coral range; for example, the extension of branching Acropora polewards has been recorded in Florida, despite an almost Caribbean-wide trend for reef deterioration (Precht and Aronson, 2004), but there are several constraints, including low genetic diversity and the limited suitable substrate at the latitudinal limits to reef growth (Riegl, 2003; Ayre and Hughes, 2004; Woodroffe et al., 2005).
The fate of the small reef islands on the rim of atolls is of special concern. Small reef islands in the Indo-Pacific formed over recent millennia during a period when regional sea level fell (Dickinson, 2004; Woodroffe and Morrison, 2001). However, the response of these islands to future sea-level rise remains uncertain, and is addressed in greater detail in Chapter 16, Section 16.4.2. It will be important to identify critical thresholds of change beyond which there may be collapse of ecological and social systems on atolls. There are limited data, little local expertise to assess the dangers, and a low level of economic activity to cover the costs of adaptation for atolls in countries such as the Maldives, Kiribati and Tuvalu (Barnett and Adger, 2003; Chapter 16, Box 16.6).
C2.2.3 Climate change and the Great Barrier Reef (Chapter 11, Box 11.3)
The Great Barrier Reef (GBR) is the world's largest continuous reef system (2,100 km long) and is a critical storehouse of Australian marine biodiversity and a breeding ground for seabirds and other marine vertebrates such as the humpback whale. Tourism associated with the GBR generated over US$4.48 billion in the 12-month period 2004/5 and provided employment for about 63,000 full-time equivalent persons (Access Economics, 2005). The two greatest threats from climate change to the GBR are (i) rising sea temperatures, which are almost certain to increase the frequency and intensity of mass coral bleaching events, and (ii) ocean acidification, which is likely to reduce the calcifying ability of key organisms such as corals. Other factors, such as droughts and more intense storms, are likely to influence reefs through physical damage and extended flood plumes (Puotinen, 2006).
Sea temperatures on the GBR have warmed by about 0.4°C over the past century (Lough, 2000). Temperatures currently typical of the northern tip of the GBR are very likely to extend to its southern end by 2040 to 2050 (SRES scenarios A1, A2) and 2070 to 2090 (SRES scenarios B1, B2) (Done et al., 2003). Temperatures only 1 °C above the long-term summer maxima already cause mass coral bleaching (loss of symbiotic algae). Corals may recover but will die under high or prolonged temperatures (2 to 3°C above long-term maxima for at least 4 weeks). The GBR has experienced eight mass bleaching events since 1979 (1980, 1982, 1987, 1992, 1994, 1998, 2002 and 2006); there are no records of events prior to 1979 (Hoegh-Guldberg, 1999). The most widespread and intense events occurred in the summers of 1998 and 2002, with about 42% and 54% of reefs affected, respectively (Done et al., 2003; Berkelmans et al., 2004). Mortality was distributed patchily, with the greatest effects on near-shore reefs, possibly exacerbated by osmotic stress caused by floodwaters in some areas (Berkelmans and Oliver, 1999). The 2002 event was followed by localised outbreaks of coral disease, with incidence of some disease-like syndromes increasing by as much as 500% over the past decade at a few sites (Willis et al.,
2004). While the impacts of coral disease on the GBR are currently minor, experiences in other parts of the world suggest that disease has the potential to be a threat to GBR reefs. Effects from thermal stress are likely to be exacerbated under future scenarios by the gradual acidification of the world's oceans, which have absorbed about 30% of the excess CO2 released to the atmosphere (Orr et al., 2005; Raven et al.,
2005). Calcification declines with decreasing carbonate ion concentrations, becoming zero at carbonate ion concentrations of approximately 200 ^mol/kg (Langdon et al., 2000; Langdon, 2002). These occur at atmospheric CO2 concentrations of approximately 500 ppm. Reduced growth due to acidic conditions is very likely to hinder reef recovery after bleaching events and will reduce the resilience of reefs to other stressors (e.g., sediment, eutrophication).
Even under a moderate warming scenario (A1T, 2°C by 2100), corals on the GBR are very likely to be exposed to regular summer temperatures that exceed the thermal thresholds observed over the past 20 years (Done et al., 2003). Annual bleaching is projected under the A1FI scenario by 2030, and under A1T by 2050 (Done et al., 2003; Wooldridge et al., 2005). Given that the recovery time from a severe bleaching-induced mortality event is at least 10 years (and may exceed 50 years for full recovery), these models suggest that reefs are likely to be dominated by non-coral organisms such as macroalgae by 2050 (Hoegh-Guldberg, 1999; Done et al., 2003). Substantial impacts on biodiversity, fishing and tourism are likely. Maintenance of hard coral cover on the GBR will require corals to increase their upper thermal tolerance limits at the same pace as the change in sea temperatures driven by climate change, i.e., about 0.1-0.5°C/decade (Donner et al., 2005). There is currently little evidence that corals have the capacity for such rapid genetic change; most of the evidence is to the contrary (Hoegh-Guldberg, 1999, 2004). Given that recovery from mortality can be potentially enhanced by reducing local stresses (water quality, fishing pressure), management initiatives such as the Reef Water Quality Protection Plan and the Representative Areas Programme (which expanded totally protected areas on the GBR from 4.6% to over 33%) represent planned adaptation options to enhance the ability of coral reefs to endure the rising pressure from rapid climate change.
C2.2.4 Impact of coral mortality on reef fisheries (Chapter 5, Box 5.4)
Coral reefs and their fisheries are subject to many stresses in addition to climate change (see Chapter 4). So far, events such as the 1998 mass coral bleaching in the Indian Ocean have not provided evidence of negative short-term bio-economic impacts for coastal reef fisheries (Spalding and Jarvis, 2002; Grandcourt and Cesar, 2003). In the longer term, there may be serious consequences for fisheries production that result from loss of coral communities and reduced structural complexity, which result in reduced fish species richness, local extinctions and loss of species within key functional groups of reef fish (Sano, 2004; Graham et al., 2006).
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