In addition to assessing WSB outbreaks using the ratio of trees that record an outbreak, the corrected chronologies were also examined to describe the integrated stand-level response to WSB outbreaks. All of the

corrected chronologies were truncated to the year 1632 and correlated to one another using Pearson correlation coefficients. Data were then transformed using a 10-year spline to reduce inter-annual variability while still maintaining high-frequency variability in the time series. All of the smoothed corrected indices MK-8776 manufacturer were grouped on the basis of their correlation coefficients and averaged into sub-regional chronologies to create outbreak histories within the larger study area. While it was not the primary objective of this study, we examined possible relationships between synchronous outbreaks and climate by comparing the sub-regional chronologies with independently reconstructed summer temperature (June–August) and May 1 snow water equivalence (SWE) anomalies

for the Tatlayoko Lake station (Table 3; Starheim et al., 2012). To facilitate comparison between the datasets, the reconstructed climatic anomalies were transformed using a 10-year spline. Synchronous WSB outbreaks were defined as periods when >5 consecutive years had index values in the lowest 75% percentile in at least 3 of the 4 sub-regional chronologies. Wavelet analysis was performed to decompose the sub-regional chronologies into time–frequency domains to identify the dominant modes of variability through time (Torrence and Compo, 1998). Wavelet analysis was performed using a continuous Morlet transformation at the 99% Nutlin-3 ic50 confidence level on the sub-regional chronologies in the R package dplR ( Bunn, 2008 and Bunn et al., 2012). The tree-ring chronologies used in this study were collected at sites found throughout the study area (Fig. 1). Fourteen archived and newly collected Douglas-fir chronologies

sites were combined to develop 11 host chronologies (Table 1). Six archived lodgepole pine chronologies and 6 archived ponderosa pine chronologies O-methylated flavonoid were combined to develop two regional non-host chronologies (Table 1). The Douglas-fir chronologies were constructed from trees found primarily in the dry-cool Fraser or the dry-cool Chilcotin BEC units, with the exception of the Fraser River and Farwell Canyon chronologies constructed from trees located in the very dry-mild BEC unit (Table 1 and Table 2). Two chronologies were located in transitional BEC units: the Bull Canyon chronology is transitional between very dry-mild and dry-cool Chilcotin; in the southeast the Chasm chronology is transitional between the very dry-warm and dry-cool Fraser (Fig. 1; Table 1 and Table 2). All the Douglas-fir sites were characterized by open forests (averaging 375 trees per hectare) where the drier stands (very dry-mild and very dry-warm) represent a transition from grassland to more continuous forest at higher elevations (dry-cool BEC units) (Steen and Coupé, 1997).