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climatic gradients and move into new areas of suit- <br />able habitat. In addition, current reserve networks, <br />many of which protect only a small, potentially <br />biased samples of environmental conditions across <br />the range of an individual species or habitat type, <br />are likely to become increasingly less representa- <br />tive under combined impacts of climate change and <br />habitat loss (Dyke 2004). <br />The negative consequences of expanding urban <br />development on species and habitats are well estab- <br />lished (e.g., Chace and Walsh 2006, Hansen et al. <br />2005, McKinney 2002). Direct habitat loss, habi- <br />tat fragmentation, and isolation can pose signifi- <br />cant threats to population viability on their own. <br />However, interactions between climate change and <br />other drivers, such as land use change, may have <br />greater impacts on biodiversity than any one driver <br />alone (Brook et al. 2008). In a survey of 248 papers <br />from the climate change literature that addressed <br />the conservation and management of biodiversity or <br />ecosystems, Felton et al. (2009) found that fewer than <br />half of the studies addressed climate change in rela- <br />tion to other anthropogenic threats. Recent reviews <br />emphasize that the lack of integration of climate <br />change impacts with other synergistic threats is likely <br />to inadequately capture future impacts on biodiver- <br />sity (Brook et al. 2008, de Chazal and Rounsevell <br />2009). For example, Warren et al. (2001) exam- <br />ined responses of 46 species of butterflies that were <br />expected to expand their ranges as a result of climate <br />warming over the last 30 years, and found that three - <br />quarters had declined as a result of habitat loss, with <br />sedentary species and habitat specialists among the <br />most adversely affected. Jetz et al. (2007) estimated <br />that globally 10 - 20% of land bird species would <br />be imperiled by climate change and land conversion <br />by 2100, but that differences in species diversity and <br />range will affect the relative influence of these driv- <br />ers. For example, they suggest that climate change <br />will be the principal driver of range contractions at <br />higher latitudes, while land conversion will be the <br />principal driver of species range contractions in the <br />tropics (Jett et al. 2007). However, their analysis did <br />not examine the potential for range shifts (in addi- <br />tion to contractions) to occur as a result of climate <br />change. <br />Projections of species range shifts under climate <br />change often assume that species distributions are <br />limited primarily by temperature without account- <br />ing for the spatial configuration of the landscape or <br />habitat. To address this issue, some authors have <br />attempted to integrate a metapopulation dynam- <br />ics framework with broader scale changes in species <br />ranges. Opdam and Washer (2004) characterize <br />range shifts as the result of extinction of (meta) popu- <br />lations at the warm range limit and colonization into <br />regions that have transitioned into suitable thermal <br />conditions at the cooler end of the range. The ability <br />of a species to shift into more suitable areas will be <br />a function of new climate conditions (e.g. tempera- <br />ture) and extreme weather conditions. However, the <br />authors suggest that some species metapopulations <br />will be unable to persist in areas where fragmenta- <br />tion has severely degraded habitat quality and patch <br />availabilty, and will likely exhibit range contractions <br />( Opdam and Washer 2004). Modeling work has <br />produced similar results, indicating that extinction <br />thresholds are lower under the combined effects of <br />habitat fragmentation and climate change (Travis <br />2003). Mclnery et al. (2007) further demonstrated <br />that the effect of habitat fragmentation on range <br />shifts may be dependent on species dispersal charac- <br />teristics and populations dynamics during periods of <br />climate change. Their model suggests that range shifts <br />are more successful in less fragmented (clumped) <br />landscapes for species with low colonization rates. <br />However, for species with especially strong disper- <br />sal and colonization abilities, fragmentation could <br />have the opposite effect, facilitating range shifts so <br />long as the availability of suitable habitat keeps pace <br />with climatic shifts on the landscape (Mclnerny et <br />al. 2007). <br />