Box 41 Ecological impacts of the European heatwave 2003

Anomalous hot and dry conditions affected Europe between June and mid-August, 2003 (Fink et al., 2004; Luterbacher et al., 2004; Schär et al., 2004). Since similarly warm summers may occur at least every second year by 2080 in a Special Report on Emissions Scenario (SRES; Nakicenovic et al, 2000) A2 world, for example (Beniston, 2004; Schär et al., 2004), effects on ecosystems observed in 2003 provide a conservative analogue of future impacts. The major effects of the 2003 heatwave on vegetation and ecosystems appear to have been through heat and drought stress, and wildfires.

Drought stress impacts on vegetation (Gobron et al., 2005; Lobo and Maisongrande, 2006) reduced gross primary production (GPP) in Europe by 30% and respiration to a lesser degree, overall resulting in a net carbon source of 0.5 PgC/yr (Ciais et al., 2005). However, vegetation responses to the heat varied along environmental gradients such as altitude, e.g., by prolonging the growing season at high elevations (Jolly et al., 2005). Some vegetation types, as monitored by remote sensing, were found to recover to a normal state by 2004 (e.g., Gobron et al., 2005), but enhanced crown damage of dominant forest trees in 2004, for example, indicates complex delayed impacts (Fischer, 2005). Freshwater ecosystems experienced prolonged depletion of oxygen in deeper layers of lakes during the heatwave (Jankowski et al., 2006), and there was a significant decline and subsequent poor recovery in species richness of molluscs in the River Saône (Mouthon and Daufresne, 2006). Taken together, this suggests quite variable resilience across ecosystems of different types, with very likely progressive impairment of ecosystem composition and function if such events increase in frequency (e.g., Lloret et al., 2004; Rebetez and Dobbertin, 2004; Jolly et al., 2005; Fuhrer et al., 2006).

High temperatures and greater dry spell durations increase vegetation flammability (e.g., Burgan et al., 1997), and during the 2003 heatwave a record-breaking incidence of spatially extensive wildfires was observed in European countries (Barbosa et al., 2003), with roughly 650,000 ha of forest burned across the continent (De Bono et al., 2004). Fire extent (area burned), although not fire incidence, was exceptional in Europe in 2003, as found for the extraordinary 2000 fire season in the USA (Brown and Hall, 2001), and noted as an increasing trend in the USA since the 1980s (Westerling et al., 2006). In Portugal, area burned was more than twice the previous extreme (1998) and four times the 1980-2004 average (Trigo et al., 2005, 2006). Over 5% of the total forest area of Portugal burned, with an economic impact exceeding €1 billion (De Bono et al., 2004).

Long-term impacts of more frequent similar events are very likely to cause changes in biome type, particularly by promoting highly flammable, shrubby vegetation that burns more frequently than less flammable vegetation types such as forests (Nunes et al., 2005), and as seen in the tendency of burned woodlands to reburn at shorter intervals (Vazquez and Moreno, 2001; Salvador et al., 2005). The conversion of vegetation structure in this way on a large enough scale may even cause accelerated climate change through losses of carbon from biospheric stocks (Cox et al., 2000). Future projections for Europe suggest significant reductions in species richness even under mean climate change conditions (Thuiller et al., 2005b), and an increased frequency of such extremes (as indicated e.g., by Schär et al., 2004) is likely to exacerbate overall biodiversity losses (Thuiller et al., 2005b).

been recognised as important drivers of past and present ecosystem change, particularly of biodiversity (Heywood and Watson, 1995; Fahrig, 2003).

Fire influences community structure by favouring species that tolerate fire or even enhance fire spread, resulting in a relationship between the relative flammability of a species and its relative abundance in a particular community (Bond and Keeley, 2005). As a result, many vegetation types are far from the maximum biomass predicted by regional climate alone (Bond et al., 2005). Geographical shifts in key species or fire may therefore cause fundamental community shifts (Brooks et al., 2004; Schumacher and Bugmann, 2006). Fire-prone vegetation types cover a total of 40% of the world's land surface (Chapin et al., 2002), and are common in tropical and subtropical regions (Bond et al., 2005), and the boreal region

(Harden et al., 2000) in particular. Intensified wildfire regimes driven at least partly by 20th century climate change (Gillett et al., 2004; Westerling et al., 2006), appear to be changing vegetation structure and composition with shifts from Picea- to Pinus--dominated communities and 75-95% reductions in tree densities observed in forest-tundra transition in eastern Canada (Lavoie and Sirois, 1998). By contrast, in Quebec, fire frequency appears to have dropped during the 20th century (Bergeron et al., 2001), a trend projected to continue (see Section 4.4.5; Bergeron et al., 2004). Across the entire North American boreal region, however, total burned area from fires increased by a factor of 2.5 between the 1960s and 1990s, while the area burned from human-ignited fires remained constant (Kasischke and Turetsky, 2006). In South-East Asia, by contrast, human activities have significantly altered fire regimes in ways that may be detrimental to the affected ecosystems (Murdiyarso and Lebel, 2007).

Drought facilitated the spread of human-caused fire in tropical regions during the 1997/98 El Niño (Randerson et al., 2005), affecting atmospheric trace gas concentrations such as CO, CH4 and H2 (Langenfelds et al., 2002; Novelli et al., 2003; Kasischke et al., 2005), and CO2 emissions (van der Werf et al., 2004) at hemispheric and global scales. Drought conditions increase Amazon forest flammability (Nepstad et al., 2004). Tropical forest fires are becoming more common (Cochrane, 2003), and have strong negative effects on Amazonian vegetation (Cochrane and Laurance, 2002; Haugaasen et al.,

2003), possibly even intensifying rainfall events (Andreae et al., 2004, but see Sections 4.4.1 and 4.4.5 on forest productivity trends).

Significant progress on globally applicable models of fire has been made since the TAR (Thonicke et al., 2001). Modelling suggests increases in wildfire impacts (see Sections 4.4.1 and 4.4.5) during the 21st century under a wide range of scenarios (e.g., Scholze et al., 2006). The implications of the regional and global importance of fire are manifold (Bond et al., 2005). Firstly, fire suppression strategies often have limited impact (Keeley, 2002; Schoennagel et al., 2004; Van Wilgen et al.,

2004), and the enhancement of vegetation flammability through more prevalent fire weather (Brown et al., 2004) and the resulting big wildfires threatens human settlements, infrastructure and livelihoods (e.g., Allen Consulting Group,

2005). Secondly, in some ecosystems, including islands, human-caused fires have transformed forests into more flammable shrublands and grasslands (Ogden et al., 1998). Thirdly, the drivers of flammability, such as ecosystem productivity, fuel accumulation and environmental fire risk conditions, are all influenced by climate change (Williams et al., 2001; see Sections 4.4.3,4.4.4 and 4.4.5).

The spatial impact of insect damage is significant and exceeds that of fire in some ecosystems, but especially in boreal forests (Logan et al., 2003). Spruce bud worm (SBW), for example, defoliated over 20 times the area burned in eastern Ontario between 1941 and 1996 (Fleming et al., 2002). Furthermore, fires tended to occur 3 to 9 years after a SBW outbreak (Fleming et al., 2002), suggesting a greater interaction between these disturbances with further warming. Disturbance by forest tent caterpillar has also increased in western Canada in the past 25 years (Timoney, 2003). In the Mediterranean region, the defoliation of Scots Pine shows a significant association with previous warm winters, implying that future climatic warming may intensify insect damage (Hodar and Zamora, 2004; see Section 4.4.5).

Invasive alien species (IAS) (Chornesky and Randall, 2003) represent a major threat to endemic or native biodiversity in terrestrial and aquatic systems (Sala et al., 2000; Scavia et al., 2002; Occhipinti-Ambrogi and Savini, 2003). Causes of biological invasions are multiple and complex (Dukes and Mooney, 1999), yet some simple models have been developed (Crawley, 1989; Deutschewitz et al., 2003; Chytry et al., 2005; Facon et al., 2006). Alien species invasions also interact with other drivers, sometimes resulting in some unexpected outcomes (Chapuis et al., 2004). Changes in biotic and/or abiotic disturbance regimes are recognised as primary drivers of IAS (Le Maitre et al., 2004), with communities often becoming more susceptible to invasion following extreme events (Smith and Knapp, 1999), such as are projected under future climate change. IAS can also change disturbance regimes through increasing vegetation flammability (Brooks et al., 2004). Overall, ongoing shifts in human-mediated disturbances, insect pests, IAS and fire regimes are very likely to be important in altering regional ecosystem structure, diversity and function (e.g., Timoney, 2003).

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