Catastrophic hydraulic failure will likely be an important mechanism contributing to large-scale tree dieback caused by increased frequency and intensity of droughts under global climate change. To compare the susceptibility of 22 temperate deciduous tree and shrub species to hydraulic failure during a record drought in the southeastern USA, we quantified leaf desiccation, native embolism, wood density, stomatal conductance and predawn and midday leaf water potential at four sites with varying drought intensities. At the two driest sites, there was widespread leaf wilting and desiccation, and most species exhibited predawn leaf water potentials of 3 MPa and 460% loss of xylem conductivity in branches. Although species with high wood density were more resistant to cavitation, they had higher levels of native embolism and greater canopy dieback than species with low wood density. This unexpected result can be explained by the failure of species with dense wood to avert a decline in water potential to dangerous levels during the drought. Leaf water potential was negatively correlated with wood density, and the relationship was strongest under conditions of severe water deficit. Species with low wood density avoided catastrophic embolism by relying on an avoidance strategy that involves partial drought deciduousness, higher sensitivity of stomata to leaf water potential and perhaps greater rooting depth. These species therefore maintained water potential at levels that ensured a greater margin of safety against embolism. These differences among species may mediate rapid shifts in species composition of temperate forests if droughts intensify due to climate change.
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Anthropogenic climate change has altered temperate forest phenology, but how these trends will play out in the future is controversial. We measured the effect of experimental warming of 0.6-5.0 °C on the phenology of a diverse suite of 11 plant species in the deciduous forest understory (Duke Forest, North Carolina, USA) in a relatively warm year (2011) and a colder year (2013). Our primary goal was to dissect how temperature affects timing of spring budburst, flowering, and autumn leaf coloring for functional groups with different growth habits, phenological niches, and xylem anatomy. Warming advanced budburst of six deciduous woody species by 5-15 days and delayed leaf coloring by 18-21 days, resulting in an extension of the growing season by as much as 20-29 days. Spring temperature accumulation was strongly correlated with budburst date, but temperature alone cannot explain the diverse budburst responses observed among plant functional types. Ring-porous trees showed a consistent temperature response pattern across years, suggesting these species are sensitive to photoperiod. Conversely, diffuse-porous species responded differently between years, suggesting winter chilling may be more important in regulating budburst. Budburst of the ring-porous Quercus alba responded nonlinearly to warming, suggesting evolutionary constraints may limit changes in phenology, and therefore productivity, in the future. Warming caused a divergence in flowering times among species in the forest community, resulting in a longer flowering season by 10-16 days. Temperature was a good predictor of flowering for only four of the seven species studied here. Observations of interannual temperature variability overpredicted flowering responses in spring-blooming species, relative to our warming experiment, and did not consistently predict even the direction of flowering shifts. Experiments that push temperatures beyond historic variation are indispensable for improving predictions of future changes in phenology.
Nutrient resorption from senescing leaves is an important mechanism of nutrient conservation in temperate deciduous forests. Resorption, however, may be curtailed by climatic events that cause rapid leaf death, such as severe drought, which has been projected to double by the year 2100 in the eastern United States. During a record drought in the southeastern US, we studied 18 common temperate winter-deciduous trees and shrubs to understand how extreme drought affects nutrient resorption of the macronutrients N, P, K, and Ca. Four species exhibited drought-induced leaf senescence and maintained higher leaf water potentials than the remaining 14 species (here called drought-evergreen species). This strategy prevented extensive leaf desiccation during the drought and successfully averted large nutrient losses caused by leaf desiccation. These four drought-deciduous species were also able to resorb N, P, and K from drought-senesced leaves, whereas drought-evergreen species did not resorb any nutrients from leaves lost to desiccation during the drought. For Oxydendrum arboreum, the species most severely affected by the drought, our results indicate that trees lost 50% more N and P due to desiccation than would have been lost from fall senescence alone. For all drought-deciduous species, resorption of N and P in fall-senesced leaves was highly proficient, whereas resorption was incomplete for drought-evergreen species. The lower seasonal nutrient losses of drought-deciduous species may give them a competitive advantage over drought-evergreen species in the years following the drought, thereby impacting species composition in temperate deciduous forests in the future.
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