The potential influence of changing climate on the persistence of salmonids of the inland west
The Earth's climate warmed steadily during the 20th century, and mean annual air temperatures are estimated to have increased by 0.6°C (Intergovernmental Panel on Climate Change, 2007). Although many cycles of warming and cooling have occurred in the past, the most recent warming period is unique in its rate and magnitude of change (Siegenthaler and others, 2005) and in its association with anthropogenic emissions of greenhouse gases (Intergovernmental Panel on Climate Change , 2007). The climate in the western United States warmed in concert with the global trend but at an accelerated rate (+0.8°C during the 20th century; Saunders and others, 2008). The region could also prove especially sensitive to future changes because the relatively small human population is growing rapidly, as are demands on limited water supplies. Regional hydrological patterns are dominated by seasonal snow accumulation at upper elevations. Most of the region is relatively dry, and both terrestrial and aquatic ecosystems are strongly constrained b y water availability (Barnett and others, 2008; Brown and others, 2008). Stream environments are dynamic and climatically extreme, and salmonid fishes are the dominant elements of the native biodiversity (McPhail and Lindsey, 1986; Waples and others, 2008). Salmonids have broad economic and ecologic importance, but a century of intensive water resource development, nonnative fish stocking, and land use has significantly reduced many populations and several taxa are now protected under the Endangered Species Act (Thurow and others, 1997; Trotter, 2008). Because salmonids require relatively pristine, cold water environments and are often isolated in headwater habitats, members of this group may be especially vulnerable to the effects of a warming climate (Keleher and Rahel, 1996; Rieman and others, 2007; Williams and others, 2009). Warming during the 20th century drove a series of environmental trends that have profound implications for many aspects of salmonid habitat, including disturbance regimes such as wildfire, and unfavorable changes to thermal and hydrologic properties of aquatic systems. Warmer air temperatures have been associated with decreased winter snow accumulations, have accelerated snowmelt, and have advanced the timing of peak runoff by several days to weeks across most of western North America (Stewart and others, 2005; Barnett and others, 2008). Less snow and earlier runoff decrease aquifer recharge, make less water available for groundwater inputs to streams, and are contributing to widespread decreases in summer low flows (Stewart and others, 2005; Rood and others, 2008; Luce and Holden 2009). Interannual variability in stream flow is increasing, as is the persistence of multi-year extreme conditions (McCabe and others, 2004; Pagano and Garen 2005). In many areas of western North America, flood risks have increased in association with warmer temperatures during the 20th century (Hamlet and Lettenmaier, 2005). Streams where midwinter temperatures are near freezing have proven especially sensitive to increased flooding because of associated transitional hydrological patterns (mixtures of rainfall and snowmelt) and propensity for occasional rain-on-snow events to rapidly melt winter snowpack and generate large floods (Hamlet and Lettenmaier, 2005). Stream temperatures in many areas are increasing (Peterson and Kitchell, 2001; Morrison and others, 2002; Bartholow, 2005; Kaushal and others, 2010), due to both air temperature increases and reduced summer flows that make streams more sensitive to warmer air temperatures (Isaak and others, 2010). In recent decades, wildfires have become more common across much of the western United States during periods of more frequent droughts (Westerling and others, 2006; Hoerling and Eischeid, 2007), and local stream temperature can increase in postfire environments (Gresswell, 1999; Dunham and others, 2007). Fire-related temperature increase within streams is commonly a transient phenomenon, lasting only until riparian vegetation has recovered (Gresswell, 1999); however, ongoing climate change could preclude recovery to higher stature, prefire vegetation types in some areas (McKenzie and others, 2004; van Mantgem and Stephenson, 2007), resulting in a loss of critical riparian shading. Additionally, when wildfires occur in steep mountain topographies, the vegetation that stabilize s soils on hillslopes is often killed and landslides become more prevalent (Gresswell, 1999). Landslides int o stream channels form debris flows composed of sediment slurries and dead trees that can scour channels to bedrock and further exacerbate stream heating, delay recovery of riparian areas, or extirpate fish populations (Gresswell, 1999; May and Gresswell, 2003; Dunham and others, 2007). Changes in stream environments will shift habitat distributions, sometimes unpredictably, in both time and space for many salmonid fishes. Water temperature fundamentally influences aquatic ecosystem health because distribution, reproduction, fitness, and survival of ectothermic organisms are inextricably linked to the thermal regime of the environment. Historically, research has focused on defining lethal thermal limits of salmonids (Eaton and others, 1995; Selong and others, 2001; Todd and others, 2008); however, water temperature is known to be important in biological processes at a variety of spatial scales and levels of biological organization (Rahel and Olden, 2008; McCullough and others, 2009). For instance, trout are affected directly by water temperature through feeding, metabolism, and growth rates, and indirectly by factors such as prey availability and species interactions (Wehrly and others, 2007; Rahel and Olden, 2008). Where cold water temperatures currently limit habitat suitability and distributions of some species (for example, at the highest and most northerly distributional extents; Nakano and others, 1996; Coleman and Fausch, 2007), a warming climate may gradually increase the quality and extent of suitable habitat. Over time, previously constrained populations are expected to expand into these new habitats and increase in number. Some evidence suggests this may already be happening in Alaska, where streams in recently deglaciated areas are being colonized by emigrants from nearby salmon and char populations (Milner and others, 2000). Unfortunately, many of the sensitive salmonid species that are often the focus of western managers are unlikely to benefit from future water temperature increases. Warmer stream temperatures will facilitate invasion by nonnative species that are broadly established in downstream areas into upstream areas where they will compete with native species (Rieman and others, 2006; Rahel and Olden, 2008; Fausch and others, 2009). In other cases, warmer stream temperatures will render thermally suitable habitats unsuitable in downstream areas and effect net losses of habitat because upstream distributions are often constrained by streams that are too small or steep (Hari and others, 2006; Isaak and others, 2010). Both scenarios are realistic for fish species like bull trout ( Salvelinus confluentus ) (Rieman and others, 2006; Rieman and others, 2007), the various subspecies of cutthroat trout ( Oncorhynchus clarkii ) (Williams and others, 2009), Gila trout ( Oncorhynchus gilae gilae ) (Kennedy and others, 2008), and Apache trout ( Oncorhynchus gilae apache ) (Rinne and Minckley, 1985; Carmichael and others, 1993). As native species are increasingly confined to smaller and more isolated habitats by a gradually warming climate, the effects of wildfires (whether related to lethal changes in water quality during a fire, channel debris flows, or chronic postfire warming ) could have greater proportional effects on remaining habitats (for example, Brown and others, 2001; Rieman and others, 2007). If these changes were accompanied by additional hydrologic alterations associated with changes to the magnitude, frequency, duration, timing, and rate of change of discharge patterns (Jager and others, 1999; Henderson and others, 2000), populations may begin to lose some of their historic resilience and become ever more susceptible to local extirpations. As dramatic and extensive as climatic and environmental trends are for salmonid habitats, global climate models (GCMs) project that many of these trends will continue and even accelerate until at least the middle of the 21st century (Intergovernmental Panel on Climate Change, 2007). Current projections suggest mean annual air temperatures will increase by an additional 1–3°C, and early indications are that climate trajectory is at the higher end of this range (Pittock, 2006; Raupach and others, 2007). Although predicted changes vary considerably, even the most conservative estimates suggest a warming rate that will be twice that observed during the 20th century. Projections for the midcentury are most certainly due to the effects of greenhouse gases already emitted or predicted in the short term, uncertainties of the effects of longer-term greenhouse gas emissions, short-term climate cycles, and process errors associated with climate models (Cox and Stephenson, 2007). Projections of changes in total precipitation are less certain than those for air temperatures, but most GCMs project relatively small changes in the Northwest, with the exception of slightly drier summer periods (Mote and others, 2008; Karl and others, 2009). In the Southwest, however, significant decreases (such as 15–30 percent ) are projected during most periods of the year, and this area is one of the few for which Intergovernmental Panel on Climate Change (2007) precipitation projections have a high level of certainty (Hoerling and Eischeid, 2007; Karl and others, 2009). Clearly, managers of native salmonids in the wester n United States should consider adjusting management strategies to accommodate a warmer and possibly drier future (Williams and others, 2009). Tools are needed to forecast where important changes may occur and how conservation efforts should be prioritized. In this Open-File Report, we document our initial efforts in this regard for 10 species and subspecies of inland trout and Montana Arctic grayling ( Thymallus arcticus ) across the western United States.