Physiological limitation at alpine treeline: relationships of threshold responses of conifers to their establishment patterns

An understanding of physiological limitations to tree establishment at alpine treeline form the basis for predicting how this climate-driven boundary will respond to climate shifts. Most research on this topic has focused on limitations related to carbon balance and growth of trees. Carbon balance could limit survival and establishment primarily through slow-acting, chronic means. We asked whether tree survival and thus establishment patterns reflect control by chronic effects in comparison to acute, threshold responses, such as survival of frost events. Seedling survivorship patterns were compared to thresholds in freezing (temperature causing leaf freezing, or freezing point, FP; and physiological response to freezing) and water status (turgor loss point, TLP; and related physiological adjustments). Subject seedlings were from forest, treeline, and alpine sites in the Alpine Treeline Warming Experiment in Colorado, and included limber and lodgepole pine (a low-elevation species), and Engelmann Spruce. Preliminary results show survival increases with seedling age, but the only corresponding increase in stress acclimation was photosynthetic resistance to freezing and TLP, not FP. Differences in survivorship among the species were not consistent with variation in FP but they generally agreed with variation in photosynthetic resistance to deep freezing and to early-season drought avoidance. Mortality of limber pine increased 35% when minimum temperatures decreased below -9C, which compares with FPs of >-8.6C, and about 1/3 of its mortality occurred during cold/wet events, particularly in the alpine. The other major correlate of mortality is midsummer drying events, as previously reported. Also in limber pine, the TLP for year-old seedlings (-2.5 MPa) corresponded with seasonal-drought mortality. In summary, we show several examples of correspondence in physiological thresholds to mortality events within a species, although the relationships are not strong. Across species, photosynthetic resistance to freezing and early-season drought avoidance related well to mortality patterns. These results are generally more supportive of the role of chronic rather than acute climate effects in broad patterns of tree seedling establishment at treeline.