Mortality in Subalpine Forests of the Sierra Nevada, California, USA: Differential Response of Pines (Pinus albicaulis and P. flexilis) to Climate Variability

Widespread forest mortality in high-elevation forests has been increasing across western North American mountains in recent years, with climate, insects, and disease the primary causes. Subalpine forests in the eastern Sierra Nevada, by contrast, have experienced far less mortality than other ranges, and mortality events have been patchy and episodic. This situation, and lack of significant effect of non-native white-pine blister rust, enable investigation of fine-scale response of two subalpine Sierran species, whitebark pine (Pinus albicaulis, PiAl) and limber pine (P. flexilis, PiFl), to climate variability. We report similarities and differences between the two major mortality events in these pines in the last 150 years: 1988-1992 for PiFl and 2006-ongoing for PiAl. In both species, the events occurred within monotypic, closed-canopy, relatively young stands (< 200 yrs PiAl, < 300 yrs in PiFl); were localized to central-eastern Sierra Nevada; and occurred at 2740-2840 m along the eastern edge of the escarpment on north/northeast aspects with slopes > 40%. Mortality patches averaged 40-80 ha in both species, with mean stand mortality of trees > 10 cm diameter 91% in PiAl and 60% in PiFl. The ultimate cause of tree death was mountain pine beetle (Dendroctonus ponderosae) in both species, with increasing 20th/21st C minimum temperatures combined with drought the pre-conditioning factors. Overall growth in the past 150 years suggests that PiFl is more drought hardy than PiAl but responds sensitively to the combined effects of drought and increasing warmth. After the 1988-1992 drought, surviving PiFl recovered growth. PiAl trees grew very poorly during that drought, and continued poor growth in the years until 2006 when the mortality event occurred in PiAl. A significant species effect is the apparent difference in levels of within-stand genetic diversity for climate factors. Differential growth between 19th C (cool, wet) and 20th/21st C (warming, drying) of PiFl trees that died versus survivors indicates that considerable within-stand genetic diversity for climate existed in PiFl. For PiFl, the late 20th C mortality event acted as strong natural selection to improve within-stand fitness for warmer and drier conditions. PiFl trees that survived the 1988-1992 drought remained healthy through subsequent droughts, including the drought that is currently causing PiAl mortality. By contrast, the PiAl stands do not appear to have contained adaptive genetic diversity for drought and warmth, and PiAl trees growth behavior over the past 150 years was similar in pattern to the PiFl trees that died. As a result, the mortality event in PiAl is creating forest openings, with unknown future stand conditions, rather than rapid within-species adaptation that occurred in PiFl.