Tree demography on Mt. Rainier: Forecasting range shifts under global warming Janneke Hille Ris Lambers, Ailene Kane, Andrew Larson & Jessica Lundquist - University of Washington Introduction Humans are changing the weather – temperatures are rising and snowfall amounts are decreasing at higher latitudes and on mountain tops. One of the greatest challenges ecologists face is forecasting how such climatic changes will affect species distributions. There is ample evidence that species’ ranges are already shifting in response to warming, and models suggest that continued migration poleward and uphill is needed for species to track areas climatically similar to those they occur in now. Forecasting the rate of these range shifts requires an understanding of how climate influences demography (growth, survival, fecundity) across species’ ranges. With its 4400 meters in elevation and a strong rainshadow, Mt. Rainier is a microcosm of climate zones and forest types found in Oregon, Washington and British Columbia. It is thus an ideal location for studies of how climate change will affect tree range shifts. Here, we use data collected from Mt. Rainier National Park to understand the relationship between elevation, climate parameters and tree performance for 4 conifers (Douglas fir, Western hemlock, Western red cedar, Pacific silver fir). We ask: 1. Does abundance, growth and survival of these 4 tree species, measured at 18 permanent forest stands, correlate with temperature, precipitation or snowfall? 2. Do annual growth rates of adult trees and juvenile seedlings, measured at one location, correlate with annually varying snow, temperature, and precipitation? Conclusions Results 1. Climate correlates with the abundance of 1 species. Although the abundance of Douglas fir, Western red cedar and Western hemlock decline with elevation (Fig 2.), only Western hemlock abundance correlates with three climate variables (Fig. 3). It is possible that these climate variables drive the abundance of this species at Mt. Rainier, but our analyses were limited by small sample size (18 stands) and correlations between the climate variables (Table 1). Figure 3. The relationship between mean annual temperature, total precipitation, and total snowfall and the basal area of Western hemlock at 18 stands across Mt. Rainier. Precipitation and snowfall are reported as mm of rain equivalent. Table 1. Correlations between climate variables at 18 stands Climate variable Elevation Precipitation Snow Temperature Elevation Precipitation 0.404 Snow 0.609 0.876 Temperature -0.701 -0.484 -0.832 2. Snow influences Western hemlock demography, and may affect other species by increasing late-season soil moisture while decreasing growing season length. 3. Tree rings show great promise for assessing impacts of global warming on population dynamics. 2. Stand-specific growth and survival rates of 1 species was correlated with average annual snowfall. Methods We use three kinds of data: 1. Growth and mortality data were collected from 18 one hectare stands within the park (Fig. 1). These stands were established in 1978, with the size and survival of each tree censused every 5 years. The stands vary widely in climatic variables and the abundance of the four focal tree species (Fig. 2). 2. Annual ring widths quantified from increment cores and seedling crosssections. Data were collected adjacent to Longmire, a climate monitoring station (Fig. 2). We collected ten increment cores and 10 seedling crosssections for 2 species (Western hemlock, Pacific silver fir), and measured ringwidths using WinDENDRO. 3. Climate averages were estimated for each stand using the Parameter-elevation Regressions on Independent Slopes Model (PRISM) climate mapping system (PRISM Group 2007). Climate data for Longmire were acquired from Natural Resource Conservation Service and the National Climate Data Center. For all trees, growth and survival depended on tree size. Thus, we fit a relationship between size and growth and size and survival (power-law and logistic, respectively) using maximum likelihood, and asked whether parameters describing size-specific growth and survival Figure 4. The relationship between snowfall and stand-specific growth parameters (from power law sizerates correlated with climate growth relationships), and size-growth relationships of two stands spanning the extremes of snowfall. variables. For all but one species, size-specific growth and survival was not correlated with stand-specific climate variables. For survival data, low sample size affected model fitting and interpretation (in many stands, fewer than 5 trees died over the 30 year sampling period). However, Western hemlock growth was correlated with snow parameters, with lower growth in plots where snowfall is higher (Fig. 4). 3. Tree ring data indicates that the annual growth of two species will be sensitive to climate change. Annual growth rates of Western hemlock and Pacific silver fir seedlings and adults at 1000 meters, where both these species are abundant, correlate with annual fluctuations in precipitation, snowfall and/or temperature (Table 2). Climate variables do not explain much of the annual variation in growth, presumably due to the low sample size (5 individuals per size class and species). Decreased snow with warming will Table 2. Coefficients and model R2’s from a linear regressions between likely positively affect Western hemlock annual growth chronologies and climate variables. Western hemlock Pacific silver fir recruitment and growth, although the Climate variable Seedling Adult Seedling Adult negative effect of increased temp0.0037 0.0001 0.0011 -0.0001 erature on adult growth may dampen Precipitation (total) -0.0014 -0.0008 0.0005 0.0052 this response. By contrast, increased Snow temperatures will positively affect -0.0029 -0.0451 0.1041 -0.0022 Temperature Pacific silver fir seedling growth. Model fit (R squared) 0.308 0.100 0.135 0.050 Figure 1. Mt. Rainier National Park (solid line), roads (dashed lines), stand locations (black dots), and site of long-term climate records (blue star). 1. Tree growth and survival of most species did not vary with climatic variables, suggesting that range shifts in response to global warming may be difficult to predict. Figure 2. Abundance of the four focal species across an altitudinal gradient. Each dot represents the species’ basal area at each of the 18 stands. For more information, email jhrl@u.washington.edu http://protist.biology.washington.edu/oikos/index.html Future Work 1. Monitoring of microclimate. To better understand how climate influences demography, we will measure temperature, soil moisture, snow depth and snow duration at each stand (Fig. 1, Fig. 5). Figure 5. Temperature sensors in trees (a) deployed using a slingshot (b) by Jessica Lundquist (c). Combining measurements of air temperature and soil temperature over time allows for quantification of snow duration (d) and and snow depth (Lundquist 2008, in press). 2. Seedling demography. The growth and survival of conifer seedlings in the Pacific Northwest may be far more sensitive to climate than are adult trees, because of shallow roots which make them sensitive to the late summer droughts common to this region, and their short stature that makes them susceptible to short growing seasons when buried by snow. Thus, we will census seedling growth and survival in each stand (Fig. 1), and compare these measures to microclimate measures (Fig. 5). 3. Quantify seed dispersal, and estimate migration rates. Forecasting range shifts with climate change requires knowledge of demographic sensitivity to altered climate AND dispersal rates. We will quantify seed dispersal, and combine estimates with demography data to forecast conifer migration rates on Mt. Rainier. Acknowledgements We thank Jerry Franklin, Steve Acker and other members of the Permanent Sample Network for graciously sharing their data, Howard Bruner for data management, Jonathan Deschamps for analyzing ring widths, and HJ Andrews LTER for providing the growth and mortality data. Funding for these data was in part provided by NSF LTER (DEB-02-18088)