Longterm Approaches to Assessing Tree Community Responses to Resource Limitation and Climate Variation
The effects of climate change on forest dynamics will be determined by tree responses at different life-stages and different scales -- from establishment to maturity and from individuals to populations. Studies incorporating local factors, such as natural enemies, competition, or tree physiology, with sufficient variation in climate are lacking. The importance of global and regional climate variation vs. local conditions and responses is poorly understood and may only be addressed with large datasets capturing sufficient environmental variation. This dissertation uses several large datasets to examine tree demographic and ecophysiological responses to light, moisture, predation, and climate in eastern temperate forests of North Carolina.
First, I use a 19-yr seed rain record from 13 forest plots in the piedmont, transition zone, and mountains to examine how climate-mediated seed maturation and density-dependent seed predation processes increase population reproductive variation in nine temperate tree species (Chapter 1). I address several hypotheses explaining interannual reproductive variation, such as resource matching, predator satiation, and pulse resource dynamics. My results indicate that (1) interannual reproductive variation increased as a result of seed maturation and seed predation processes, (2) seed maturation rates increased under warm, wet conditions, and (3) seed predation rates exhibited negative and positive density-dependence, depending of tree species and type of seed predator (specialist insects vs. generalist vertebrates). Because positive density-dependent seed predation dampened and negative density-dependent seed predation amplified the effects of climate-mediated maturation on reproductive variation, this study showed evaluations of tree reproduction need to incorporate both climate and seed predation.
Next, I use an 11-yr record of annual tree seedling growth and survival in 20 tree species planted in the piedmont and mountains to quantify individual tree seedling growth and survival responses to spatial variation in resources and temporal variation in climate (Chapter 2). First, I tested whether height-mediated growth provides an advantage to large individuals in all environments by amplifying responses to light and moisture or only when those resources were plentiful. Second, I tested whether allometric and survival responses differed among species based on life-history strategies. Individual height amplified tree seedling growth. However, some species exhibited amplification at moderate to high resource levels as well as depression of growth in large individuals growing in low light and moisture environments. Shade intolerant species exhibited an increasing ratio of height to diameter growth and increasing survival probability with both increasing light and moisture resources. Conversely, shade tolerant species exhibited decreasing height to diameter ratio with increasing light, possibly because of biomass allocation toward acquisition of limiting light resources. Despite relative small effects of drought and winter temperature of tree seedling demography, the results of this study indicate that individual tree seedlings sensitive to light and moisture environments, such as large seedlings and seedlings of shade intolerant species, growing in shaded or xeric sites may be particularly vulnerable to climate induced mortality.
Finally, I examine interannual and interspecific variation in canopy conductance using four years of environmental (vapor pressure deficit, above canopy light, and soil moisture) and stem sap flux data from heat dissipation probes for six co-occurring tree species. I developed a state-space modeling framework for predicting canopy conductance and transpiration which incorporates uncertainty in canopy and observation uncertainty. This approach is used to evaluate the degree to which co-occur deciduous tree species exhibited drought tolerating and drought avoiding canopy responses and whether these patterns were maintained in the face of interannual variation in environmental drivers. Comparisons of canopy conductance responses to environmental forcing across species and years highlighted the importance of tree sensitivity to moisture limitation, both in terms of high vapor pressure deficit and low soil moisture, and tree hydraulic characteristics within diverse forest communities. The state-space model produced similar parameter estimates to the more traditional boundary line analysis, performed well in terms of in-sample and out-of-sample prediction of sap flux observations, and provided for coherent incorporation of parameter, process, and observation errors in predicting missing data (i.e., gap-filling), canopy conductance, and transpiration.
Much needs to be learned about forest community responses to climate change, however these responses depend on local growing conditions (light and moisture), the life-stage being examined (seedlings, juveniles, or mature trees), and the scale of inference (individuals, canopies, or populations). Because climate change will not occur in isolation from other factors, such as stand age or disturbance, studies must characterize tree responses across multidimensional gradients in growing conditions. This dissertation addresses these challenges using large demographic and ecophysiological datasets well-suited for global change research.
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