Mortality and climatic/environmental correlates of Widdringtonia cedargensis trees in the Cederberg, South Africa

Background: Conifer populations appear disproportionately threatened by global change. Most examples are, however, drawn from the northern hemisphere and long-term rates of population decline are not well documented as historical data are often lacking. We use a large and long-term (1931-2013) repeat photography dataset together with environmental data and fire records to account for the decline of the critically endangered Widdringtonia cedarbergensis. Eighty-seven historical and repeat photo-pairs were analysed to establish 20th century changes in W. cedarbergensis demography. A generalized linear mixed-effects model was fitted to determine the relative importance of environmental factors and fire-return interval on mortality for the species. Results: From an initial total of 1313 live trees in historical photographs, 74% had died and only 44 (3.4%) had recruited in the repeat photographs, leaving 387 live individuals. Juveniles (mature adults) had decreased (increased) from 27% (73%) to 8% (92%) over the intervening period. Our model demonstrates that mortality is related to greater fire frequency, higher temperatures, lower elevations, less rocky habitats and aspect (i.e. east-facing slopes had the least mortality). Conclusions: Our results show that W. cedarbergensis populations have declined significantly over the recorded period, with a pronounced decline in the last 30 years. Individuals that established in open habitats at lower, hotter elevations and experienced a greater fire frequency appear to be more vulnerable to mortality than individuals growing within protected, rocky environments at higher, cooler locations with less frequent fires. Climate models predict increasing temperatures for our study area (and likely increases in wildfires). If these predictions are realised, further declines in the species can be expected. Urgent management interventions, including seedling out-planting in fire-protected high elevation sites, reducing fire frequency in higher elevation populations, and assisted migration, should be considered.

(Table 1) Tree age, post fire stand age, and soil organic matter thickness in the western Quebec black spruce boreal ecosystem

The observed long-term decrease in the regional fire activity of Eastern Canada results in excessive accumulation of organic layer on the forest floor of coniferous forests, which may affect climate-growth relationships in canopy trees. To test this hypothesis, we related tree-ring chronologies of black spruce (Picea mariana (Mill.) B.S.P.) to soil organic layer (SOL) depth at the stand scale in the lowland forests of Quebec's Clay Belt. Late-winter and early-spring temperatures and temperature at the end of the previous year's growing season were the major monthly level environmental controls of spruce growth. The effect of SOL on climate-growth relationships was moderate and reversed the association between tree growth and summer aridity from a negative to a positive relationship: trees growing on thin organic layers were thus negatively affected by drought, whereas it was the opposite for sites with deep (>20-30 cm) organic layers. This indicates the development of wetter conditions on sites with thicker SOL. Deep SOL were also associated with an increased frequency of negative growth anomalies (pointer years) in tree-ring chronologies. Our results emphasize the presence of nonlinear growth responses to SOL accumulation, suggesting 20-30 cm as a provisional threshold with respect to the effects of SOL on the climate-growth relationship. Given the current climatic conditions characterized by generally low-fire activity and a trend toward accumulation of SOL, the importance of SOL effects in the black spruce ecosystem is expected to increase in the future.