Western pine beetle voltinism in a changing California climate

Recent hot droughts in California resulted in ponderosa pine (Pinus ponderosa) mortality attributed to drought and western pine beetle (WPB, Dendroctonus brevicomis). While drought alone can cause tree death, direct warming effects on WPB are a contributing factor. Research on WPB generation timing (voltinism), however, is lacking.We monitored WPB tree attacks and adult emergence timing at two California sites and developed a degree-day model from field-observed data. Historical, contemporary, and future temperatures for several California sites were used with the model to examine trends in WPB voltinism.Field data showed a single summer and an overwinter generation at a northern California site. As summer temperatures increased beyond 1900-1980 averages, the predicted number of full and partial WPB generations by 2021 had increased from similar to 2 annual (one summer and one overwinter) generations historically to similar to 2.3 at two northern California sites and from similar to 2.3 to similar to 3.2 at two warmer California sites.Historical and contemporary data suggest winter warming was not sufficient for an additional generation overwinter. Instead, increases in generations were driven by summer and fall temperatures.Unconstrained increases in the number of future annual generations will be limited by complex, but not well understood, WPB thermal adaptations. Increased knowledge of temperature-driven WPB population growth will improve forest vegetation models aimed at predicting ponderosa pine mortality in a changing climate.