Box 44 Coral reefs endangered by climate change

Reefs are habitat for about a quarter of 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 (Chapter 6; Hoegh-Guldberg et al., 2000; Cesar et al., 2003; Willig et al., 2003; Hoegh-Guldberg, 2004, 2005) and, though 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 (Chapter 6, Box 6.1; Reynaud et al., 2003; McNeil et al., 2004; McWilliams et al., 2005) leading to bleaching (loss of algal symbionts - Chapter 6, Box 6.1). 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 (Chapter 6, Box 6.1). Annual or bi-annual exceedance of 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 demise of some reefs to 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 (Chapter 6, Section 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 subtropical 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 changes through changes to precipitation and river flow; Chapter 6, Box 6.1; Chapter 11, Box 11.3; Chapter 16)) add multiple impacts on coral reefs (Chapter 16, Box 16.2), 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).

Projections of ocean biological response to climate warming by 2050 show contraction of the highly productive marginal sea-ice biome by 42% and 17% in Northern and Southern Hemispheres (Sarmiento et al., 2004b; see also Meehl et al., 2007; Christensen et al., 2007). The sea-ice biome accounts for a large proportion of primary production in polar waters and supports a substantial food web. As timing of the spring phytoplankton bloom is linked to the sea-ice edge, loss of sea ice (Walsh and Timlin, 2003) and large reductions of the total primary production in the marginal sea-ice biome in the Northern Hemisphere (Behrenfeld and Falkowski, 1997; Marra et al., 2003) would have strong effects, for example, on the productivity of the Bering Sea (Stabeno et al., 2001). Reductions in winter sea-ice will affect the reproduction, growth and development of fish, krill, and their predators, including seals and seal-dependent polar bears (e.g., Barber and Iacozza, 2004; Box 4.3), leading to further changes in abundance and distribution of marine species (Chapter 15, Section 15.4.3). An expansion by 4.0% (Northern Hemisphere) and 9.4% (Southern), and of the sub-polar gyre biome by 16% (Northern) and 7% (Southern), has been projected for the permanently stratified sub-tropical gyre biome with its low productivity. This effect has now been observed in the North Pacific and Atlantic (McClain et al., 2004; Sarmiento et al., 2004b). A contraction by 11% of the seasonally stratified sub-tropical gyre is also projected in both hemispheres by 2050 due to climate warming. These changes are likely to have significant impacts on marine ecosystem productivity globally, with uncertainties in projections of NPP using six mainly IS92a-based scenarios narrowing to an increase of between 0.7% and 8.1% by mid-century (ATg!obal~1.5-3°C).

Changes to planktonic and benthic community composition and productivity have been observed in the North Sea since 1955 (Clark and Frid, 2001) and since the mid-1980s may have reduced the survival of young cod (Beaugrand et al., 2003). Large shifts in pelagic biodiversity (Beaugrand et al., 2002) and in fish community composition have been seen (Genner et al., 2004; Perry et al., 2005). Changes in seasonality or recurrence of hydrographic events or productive periods could be affected by trophic links to many marine populations, including exploited or cultured populations (Stenseth et al., 2002, 2003; Platt et al., 2003; Llope et al., 2006). Elevated temperatures have increased mortality of winter flounder eggs and larvae (Keller and Klein-Macphee, 2000) and have led to later spawning migrations (Sims et al., 2004). A 2°C rise in sea surface temperature (SST) would result in removal of Antarctic bivalves and limpets from the Southern Ocean (Peck et al., 2004). Tuna populations may spread towards presently temperate regions, based on predicted warming of surface water and increasing primary production at mid- and high latitudes (Loukos et al., 2003).

Marine mammals, birds, cetaceans and pinnipeds (seals, sea lions and walruses), which feed mainly on plankton, fish and squid, are vulnerable to climate change-driven changes in prey distribution, abundance and community composition in response to climatic factors (Learmonth et al., 2006). Changing water temperature also has an effect on the reproduction of cetaceans and pinnipeds, indirectly through prey abundance, either through extending the time between individual breeding attempts, or by reducing breeding condition of the mother (Whitehead, 1997). Current extreme climatic events provide an indication of potential future effects. For example, the warm-water phase of ENSO is associated with large-scale changes in plankton abundance and associated impacts on food webs (Hays et al.,

2005), and changes to behaviour (Lusseau et al., 2004), sex ratio (Vergani et al., 2004) and feeding and diet (Piatkowski et al.,

2002) of marine mammals.

Melting Arctic ice-sheets will reduce ocean salinities (IPCC, 2001), causing species-specific shifts in the distribution and biomass of major constituents of Arctic food webs, including poleward shifts in communities and the potential loss of some polar species (such as the narwhal, Monodon monoceros). Migratory whales (e.g., grey whale, Eschrichtius robustus), that spend summer in Arctic feeding grounds, are likely to experience disruptions in their food sources (Learmonth et al.,

2006). Nesting biology of sea turtles is strongly affected by temperature, both in timing and in the determination of the sex ratio of hatchlings (Hays et al., 2003), but implications for population size are unknown. A predicted sea-level rise of 0.5 m will eliminate up to 32% of sea-turtle nesting beaches in the Caribbean (Fish et al., 2005).

Surface ocean pH has decreased by 0.1 unit due to absorption of anthropogenic CO2 emissions (equivalent to a 30% increase in hydrogen ion concentration) and is predicted to decrease by up to a further 0.3-0.4 units by 2100 (Caldeira and Wickett,

2003). This may impact a wide range of organisms and ecosystems (e.g., coral reefs, Box 4.4, reviewed by Raven et al., 2005), including juvenile planktonic, as well as adult, forms of benthic calcifying organisms (e.g., echinoderms, gastropods and shellfish), and will affect their recruitment (reviewed by Turley et al., 2006). Polar and sub-polar surface waters and the Southern Ocean will be aragonite under-saturated by 2100 (Orr et al.,

2005) and Arctic waters will be similarly threatened (Haugan et al., 2006). Organisms using aragonite to make their shells (e.g., pteropods) will be at risk and this will threaten ecosystems such as the Southern and Arctic Oceans in which they play a dominant role in the food web and carbon cycling (Orr et al., 2005; Haugan et al., 2006).

Cold-water coral ecosystems exist in almost all the world's oceans and their aerial coverage could equal or exceed that of warm-water coral reefs (Freiwald et al., 2004; Guinotte et al.,

2006). They harbour a distinct and rich ecosystem, provide habitats and nursery grounds for a variety of species, including commercial fish and numerous new species previously thought to be extinct (Raven et al., 2005). These geologically ancient, long-lived, slow-growing and fragile reefs will suffer reduced calcification rates and, as the aragonite saturation horizon moves towards the ocean surface, large parts of the oceans will cease to support them by 2100 (Feely et al., 2004; Orr et al., 2005; Raven et al., 2005; Guinotte et al., 2006). Since cold-water corals do not have symbiotic algae but depend on extracting food particles sinking from surface waters or carried by ocean currents, they are also vulnerable to changes to ocean currents, primary productivity and flux of food particles (Guinotte et al., 2006). Warm-water coral reefs are also sensitive to multiple impacts including increased SST and decreasing aragonite concentrations within this century (Box 4.4).

4.4.10 Cross-biome impacts

This section highlights issues that cut across biomes, such as large-scale geographical shifts of vegetation (Figure 4.3) or animal migration patterns (e.g., Box 4.3; Box 4.5), and changes in land use and aquatic systems.

Biome shifts

Boreal forest and Arctic tundra ecosystems are projected generally to show increased growth due to longer and warmer growing seasons (Lucht et al., 2002; Figure 4.3). Woody boreal vegetation is expected to spread into tundra at higher latitudes and higher elevations (Grace et al., 2002; Kaplan et al., 2003; Gerber et al., 2004). At the southern ecotone (see Glossary) with continental grasslands, a contraction of boreal forest is projected due to increased impacts of drought, insects and fires (Bachelet et al., 2001; Scholze et al., 2006), together with a lower rate of sapling survival (Hogg and Schwarz, 1997). Drought stress could partially be counteracted by concurrent CO2-induced enhanced water-use efficiency (Gerten et al., 2005), small regional increases in precipitation, and an increased depth of permafrost thawing. It is uncertain whether peak summer heat stress on boreal tree species could cause regional transitions to grassland where the continental winter climate remains too cold for temperate forest species to succeed (Gerber et al., 2004; Lucht et al., 2006). In temperate forests, milder winters may reduce winter hardening in trees, increasing their vulnerability to frost (Hanninen et al., 2001; Hanninen, 2006).

Vegetation change in the lower to mid latitudes is uncertain because transitions between tropical desert and woody vegetation types are difficult to forecast. Climate models disagree in pattern and magnitude of projected changes in atmospheric circulation and climate variability, particularly for precipitation (e.g., with respect to the Indian and West African monsoons). For the Sahel and other semi-arid regions, increasing drought is predicted by some models (Held et al., 2005), while increased water-use efficiency is projected to cause more greening (Figure 4.3), though potentially associated with more frequent fires, in others (Bachelet et al., 2003; Woodward and Lomas, 2004b; Ni et al., 2006; Schaphoff et al., 2006). In savannas, woody encroachment is projected to be a consequence of enhanced water-use efficiency and increased precipitation in some regions (Bachelet et al., 2001; Lucht et al., 2006; Ni et al., 2006; Schaphoff et al., 2006; Section 4.4.3; Figure 4.3). The moderate drying, including desert amelioration, as projected in southern Africa, the Sahel region, central Australia, the Arabian Peninsula and parts of central Asia (Figure 4.3) may be due to a positive impact of rising atmospheric CO2, as noted in eastern Namibia through sensitivity analysis (Thuiller et al., 2006b).

A general increase of deciduous at the expense of evergreen vegetation is predicted at all latitudes, although the forests in both the eastern USA and eastern Asia appear to be sensitive to drought stress and decline under some scenarios (Bachelet et al., 2001; Gerten et al., 2005; Lucht et al., 2006; Scholze et al., 2006). Tropical ecosystems are expected to change, particularly in the Amazon, where a subset of GCMs shows strong to moderate reductions in precipitation with the consequence of transitions of evergreen tropical forest to rain-green forest or grasslands (Cox et al., 2004; Cramer et al., 2004; Woodward and Lomas, 2004b). However, representations of tropical succession remain underdeveloped in current models. The global land biosphere is projected by some models to lose carbon beyond temperature increases of 3°C (Gerber et al., 2004), mainly from temperate and boreal soils, with vegetation carbon declining beyond temperature increases above 5°C (Gerber et al., 2004). Carbon sinks persist mainly in the Arctic and in savanna grasslands (Woodward and Lomas, 2004b; Schaphoff et al., 2006). However, there is large variability between the projections of different vegetation (Cramer et al., 2001) and climate (Schaphoff et al., 2006) models for a given emissions scenario.

Migration patterns

Vagile (see Glossary) animals such as polar bears (sea-ice biome, tundra; Box 4.3) and in particular migratory animals (tundra, wetlands, lakes, tropical forests, savannas, etc.; Box 4.5) respond to impacts both within and across biomes. Many species breed in one area then move to another to spend the non-breeding season (Robinson et al., 2005). Many migratory species may be more vulnerable to climate change than resident species (Price and Root, 2005). As migratory species often move annually in response to seasonal climate changes, their behaviour, including migratory routes, is sensitive to climate. Numerous studies have found that many of these species are arriving earlier (Chapter 1 and e.g., Root et al., 2003). Changes in the timing of biological events are of particular concern because of a potential disconnect between migrants and their food resources if the phenology of each advances at different rates (Inouye et al., 2000; Root et al., 2003; Visser et al., 2004). The potential impact of climate change on migratory birds has been especially well studied (Box 4.5).

Land use

The relative importance of key drivers on ecosystem change varies across regions and biomes (Sala et al., 2000; Sala, 2005). Several global studies suggest that at least until 2050 land-use change will be the dominant driver of terrestrial biodiversity loss in human-dominated regions (Sala et al., 2000; UNEP, 2002; Gaston et al., 2003; Jenkins, 2003; Scharlemann et al., 2004; Sala, 2005). Conversely, climate change is likely to dominate where human interventions are limited, such as in the tundra, boreal, cool conifer forests, deserts and savanna biomes (Sala et al., 2000; Duraiappah et al., 2005). Assessment of impacts on biodiversity differ if other drivers than climate change are taken into account (Thomas et al., 2004a; Sala, 2005; Malcolm et al., 2006). Interactions among these drivers may mitigate or exacerbate the overall effects of climate change (Opdam and Wascher, 2004). The effects of land-use change on species through landscape fragmentation at the regional scale may further exacerbate impacts from climate change (Holman et al., 2005a; Del Barrio et al., 2006; Harrison et al., 2006; Rounsevell et al., 2006).

Global land-use change studies project a significant reduction in native vegetation (mostly forest) in non-industrialised countries and arid regions due to expansion of agricultural or urban land use driven principally by population growth,

Figure 4.3. Projected appreciable changes in terrestrial ecosystems by 2100 relative to 2000 as simulated by DGVM LPJ (Sitch etal., 2003; Gerten et al., 2004) for two SRES emissions scenarios (Nakicenovic et al., 2000) forcing two climate models: (a) HadCM3 A2, (b) ECHAM5 B1 (Lucht et al., 2006; Schaphoff et al., 2006). Changes are considered appreciable and are only shown if they exceed 20% of the area of a simulated grid cell (see Figure 4.2 for further explanations).

Figure 4.3. Projected appreciable changes in terrestrial ecosystems by 2100 relative to 2000 as simulated by DGVM LPJ (Sitch etal., 2003; Gerten et al., 2004) for two SRES emissions scenarios (Nakicenovic et al., 2000) forcing two climate models: (a) HadCM3 A2, (b) ECHAM5 B1 (Lucht et al., 2006; Schaphoff et al., 2006). Changes are considered appreciable and are only shown if they exceed 20% of the area of a simulated grid cell (see Figure 4.2 for further explanations).

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