A multi-species synthesis of physiological mechanisms in drought-induced tree mortality

Widespread tree mortality associated with drought 92 has been observed on all forested continents, and global change is expected to exacerbate vegetation vulnerability. Forest mortality has implications for future biosphere-atmosphere interactions of carbon, water, and energy balance, and is poorly represented in dynamic vegetation models. Reducing uncertainty requires improved mortality projections founded on robust physiological processes. However, the proposed mechanisms of drought-induced mortality, including hydraulic failure and carbon starvation, are unresolved. A growing number of empirical studies have investigated these mechanisms, but data have not been consistently analyzed across species and biomes using a standardized physiological framework. Here we show that xylem hydraulic failure was ubiquitous across multiple tree taxa at drought induced mortality. All species assessed had 60% or higher loss of xylem hydraulic conductivity, consistent with proposed theoretical and modelled survival thresholds. We found diverse responses in non-structural carbohydrate reserves at mortality, indicating that evidence supporting carbon starvation was not universal. Reduced non-structural carbohydrates were more common for gymnosperms than angiosperms, associated with xylem hydraulic vulnerability, and may have a role in reducing hydraulic function. Our finding that hydraulic failure at drought-induced mortality was persistent across species indicates that substantial improvement in vegetation modelling can be achieved using thresholds in hydraulic function

A multi-species synthesis of physiological mechanisms in drought-induced tree mortality

Widespread tree mortality associated with drought 92 has been observed on all forested continents, and global change is expected to exacerbate vegetation vulnerability. Forest mortality has implications for future biosphere-atmosphere interactions of carbon, water, and energy balance, and is poorly represented in dynamic vegetation models. Reducing uncertainty requires improved mortality projections founded on robust physiological processes. However, the proposed mechanisms of drought-induced mortality, including hydraulic failure and carbon starvation, are unresolved. A growing number of empirical studies have investigated these mechanisms, but data have not been consistently analyzed across species and biomes using a standardized physiological framework. Here we show that xylem hydraulic failure was ubiquitous across multiple tree taxa at drought induced mortality. All species assessed had 60% or higher loss of xylem hydraulic conductivity, consistent with proposed theoretical and modelled survival thresholds. We found diverse responses in non-structural carbohydrate reserves at mortality, indicating that evidence supporting carbon starvation was not universal. Reduced non-structural carbohydrates were more common for gymnosperms than angiosperms, associated with xylem hydraulic vulnerability, and may have a role in reducing hydraulic function. Our finding that hydraulic failure at drought-induced mortality was persistent across species indicates that substantial improvement in vegetation modelling can be achieved using thresholds in hydraulic function

Guidelines and considerations for designing field experiments simulating precipitation extremes in forest ecosystems

1. Precipitation regimes are changing in response to climate change, yet understanding of how forest ecosystems respond to extreme droughts and pluvials remains incomplete. As future precipitation extremes will likely fall outside the range of historical variability, precipitation manipulation experiments (PMEs) are critical to advancing knowledge about potential ecosystem responses. However, few PMEs have been conducted in forests compared to short‐statured ecosystems, and forest PMEs have unique design requirements and constraints. Moreover, past forest PMEs have lacked coordination, limiting cross‐site comparisons. Here, we review and synthesize approaches, challenges, and opportunities for conducting PMEs in forests, with the goal of guiding design decisions, while maximizing the potential for coordination. 2. We reviewed 63 forest PMEs at 70 sites world‐wide. Workshops, meetings, and communications with experimentalists were used to generate and build consensus around approaches for addressing the key challenges and enhancing coordination. 3. Past forest PMEs employed a variety of study designs related to treatment level, replication, plot and infrastructure characteristics, and measurement approaches. Important considerations for establishing new forest PMEs include: selecting appropriate treatment levels to reach ecological thresholds; balancing cost, logistical complexity, and effectiveness in infrastructure design; and preventing unintended water subsidies. Response variables in forest PMEs were organized into three broad tiers reflecting increasing complexity and resource intensiveness, with the first tier representing a recommended core set of common measurements. 4. Differences in site conditions combined with unique research questions of experimentalists necessitate careful adaptation of guidelines for forest PMEs to balance local objectives with coordination among experiments. We advocate adoption of a common framework for coordinating forest PME design to enhance cross‐site comparability and advance fundamental knowledge about the response and sensitivity of diverse forest ecosystems to precipitation extremes.New Hampshire Agricultural Experiment Station, Grant/Award Number: NH00071-M; Northern States Research Cooperative, Grant/Award Number: 14-DG-11242307- 142; National Science Foundation Long-Term Ecological Research, Grant/Award Number: 1637685; USDA Forest Service; University of New Hampshire; NASA, Grant/Award Number: NNX14AD31G; USDA National Institute of Food and Agriculture McIntire- Stennis Project, Grant/Award Number: NH00071-M; U.S. Department of Energy; Office of Science’s Terrestrial Ecosystem Science program; Pacific Northwest National Labs’ LDRD program; MSCA-IF 2015; EU-Horizon2020 program; NSF’s Research Coordination Network Progra

Resource limitation, tolerance,and the future of ecological plant classification

Throughout the evolutionary history of plants, drought, shade, and scarcity of nutrients have structured ecosystems and communities globally. Humans have begun to drastically alter the prevalence of these environmental factors with untold consequences for plant communities and ecosystems worldwide. Given limitations in using organ-level traits to predict ecological performance of species, recent advances using tolerances of low resource availability as plant functional traits are revealing the often hidden roles these factors have in structuring communities and are becoming central to classifying plants ecologically. For example, measuring the physiological drought tolerance of plants has increased the predictability of differences among species in their ability to survive drought as well as the distribution of species within and among ecosystems. Quantifying the shade tolerance of species has improved our understanding of local and regional species diversity and how species have sorted within and among regions. As the stresses on ecosystems continue to shift, coordinated studies of whole-plant growth centered on tolerance of low resource availability will be central in predicting future ecosystem functioning and biodiversity. This will require efforts that quantify tolerances for large numbers of species and develop bioinformatic and other techniques for comparing large number of species

Plant wax and carbon isotope response to heat and drought in the conifer Juniperus monosperma

Plant waxes, including n-alkanes, are commonly used for a wide range of paleo-applications. Several common traits of n-alkanes that are used as paleo-proxies include chain length distribution and average chain length (ACL), as well as plant wax carbon and hydrogen isotopic compositions. The effect of climate on plant wax traits has been the subject of many studies, but a common challenge with modern calibrations is disentangling the effects of species (genetic), temperature, and precipitation from one another. Here, we explore the effect of temperature and drought, independently and combined, on plant wax composition of the species Juniper monosperma in a large ecosystem-scale field manipulation experiment. We find that n-alkane concentrations significantly increase with temperature, but other parameters (including ACL) are not affected. These results support physiological studies that identify n-alkanes as an important barrier to water loss within the plant cuticle. Combined with prior studies, it appears that changes in ACL within sediments are likely controlled by changes in species composition rather than directly by changes in climate. We find little variation in the carbon isotopic composition (δ13C) of n-alkanes across the treatments whereas bulk leaf δ13C values are higher in the heat and drought treatment. Because leaf δ13C values represent a weighted C assimilation signal, these values reflect differences in leaf gas exchange among treatments, whereas the n-alkanes are synthesized when water availability is higher and differences among treatments are not significant enough to influence their values. These results have important implications for using n-alkane traits, including ACL and δ13C values, for paleoenvironmental reconstructions

Climate change risks to global forest health: Emergence of unexpected events of elevated tree mortality worldwide

Recent observations of elevated tree mortality following climate extremes, like heat and drought, raise concerns about climate change risks to global forest health. We currently lack both sufficient data and understanding to identify whether these observations represent a global trend toward increasing tree mortality. Here, we document events of sudden and unexpected elevated tree mortality following heat and drought events in ecosystems that previously were considered tolerant or not at risk of exposure. These events underscore the fact that climate change may affect forests with unexpected force in the future. We use the events as examples to highlight current difficulties and challenges for realistically predicting such tree mortality events and the uncertainties about future forest condition. Advances in remote sensing technology and greater availably of high-resolution data, from both field assessments and satellites, are needed to improve both understanding and prediction of forest responses to future climate change