Darin J. Law

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

Widespread tree mortality associated with drought 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 analysed 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.The mechanisms underlying drought-induced tree mortality are not fully resolved. Here, the authors show that, across multiple tree species, loss of xylem conductivity above 60% is associated with mortality, while carbon starvation is not universal.

The critical amplifying role of increasing atmospheric moisture demand on tree mortality and associated regional die-off

Drought-induced tree mortality, including large-scale die-off events and increases in background rates of mortality, is a global phenomenon (Allen et al., 2010) that can directly impact numerous earth system properties and ecosystem goods and services (Adams et al., 2010; Breshears et al., 2011; Anderegg et al., 2013). Tree mortality is particularly of concern because of the likelihood that it will increase in frequency and extent with climate change (McDowell et al., 2008, 2011; Adams et al., 2009; McDowell, 2011; Williams et al., 2013). Recent plant science advances related to drought have focused on understanding the physiological mechanisms that not only affect plant growth and associated carbon metabolism, but also the more challenging issue of predicting plant mortality thresholds (McDowell et al., 2013). Although some advances related to mechanisms of mortality have been made and have increased emphasis on interrelationships between carbon metabolism and plant hydraulics (McDowell et al., 2011), notably few studies have specifically evaluated effects of increasing atmospheric demand for moisture (i.e., vapour pressure deficit; VPD) on rates of tree death. In this opinion article we highlight the importance of considering the key risks of future large-scale tree die-off and other mortality events arising from increased VPD. Here we focus on mortality of trees, but our point about the importance of VPD is also relevant to other vascular plants.

Toward accounting for ecoclimate teleconnections: intra- and inter-continental consequences of altered energy balance after vegetation change

ContextVegetation is projected to continue to undergo major structural changes in coming decades due to land conversion and climate change, including widespread forest die-offs. These vegetation changes are important not only for their local or regional climatic effects, but also because they can affect climate and subsequently vegetation in other regions or continents through “ecoclimate teleconnections”.ObjectivesWe propose that ecoclimate teleconnections are a fundamental link among regions within and across continents, and are central to advancing large-scale macrosystems ecology.Methods and resultsWe illustrate potential ecoclimate teleconnections in a bounding simulation that assumes complete tree cover loss in western North America due to tree die-off, and which predicts subsequent drying and reduced net primary productivity in other areas of North America, the Amazon and elsewhere. Central to accurately modeling such ecoclimate teleconnections is characterizing how vegetation change alters albedo and other components of the land-surface energy balance and then scales up to impact the climate system. We introduce a framework for rapid field-based characterization of vegetation structure and energy balance to help address this challenge.ConclusionsEcoclimate teleconnections are likely a fundamental aspect of macrosystems ecology needed to account for alterations to large-scale atmospheric-ecological couplings in response to vegetation change, including deforestation, afforestation and die-off.

Synergistic Ecoclimate Teleconnections from Forest Loss in Different Regions Structure Global Ecological Responses

Forest loss in hotspots around the world impacts not only local climate where loss occurs, but also influences climate and vegetation in remote parts of the globe through ecoclimate teleconnections. The magnitude and mechanism of remote impacts likely depends on the location and distribution of forest loss hotspots, but the nature of these dependencies has not been investigated. We use global climate model simulations to estimate the distribution of ecologically-relevant climate changes resulting from forest loss in two hotspot regions: western North America (wNA), which is experiencing accelerated dieoff, and the Amazon basin, which is subject to high rates of deforestation. The remote climatic and ecological net effects of simultaneous forest loss in both regions differed from the combined effects of loss from the two regions simulated separately, as evident in three impacted areas. Eastern South American Gross Primary Productivity (GPP) increased due to changes in seasonal rainfall associated with Amazon forest loss and changes in temperature related to wNA forest loss. Eurasia’s GPP declined with wNA forest loss due to cooling temperatures increasing soil ice volume. Southeastern North American productivity increased with simultaneous forest loss, but declined with only wNA forest loss due to changes in VPD. Our results illustrate the need for a new generation of local-to-global scale analyses to identify potential ecoclimate teleconnections, their underlying mechanisms, and most importantly, their synergistic interactions, to predict the responses to increasing forest loss under future land use change and climate change.

Rangeland Responses to Predicted Increases in Drought Extremity

On the Ground Rangeland managers actively focus on the potential to induce a shift in a site to an alternative state, but predicted changes in climate, particularly the likelihood of more extreme drought, necessitate reevaluating risks for alternative states. Rangelands will differ in their susceptibility to undergo state changes due to climate change in general and for droughts of the future, in particular, which may be hotter. Trees, shrubs, and grasses are expected to differ in their sensitivity to drought, with trees likely being most sensitive; this affects the likelihood for state changes in grasslands, shrublands, woodlands, and savannas. Considering these differences can help rangeland managers deal with the challenges of increasing drought that is forecast to occur with climate change.

Subcontinental heat wave triggers terrestrial and marine, multi-taxa responses

Heat waves have profoundly impacted biota globally over the past decade, especially where their ecological impacts are rapid, diverse, and broad-scale. Although usually considered in isolation for either terrestrial or marine ecosystems, heat waves can straddle ecosystems of both types at subcontinental scales, potentially impacting larger areas and taxonomic breadth than previously envisioned. Using climatic and multi-species demographic data collected in Western Australia, we show that a massive heat wave event straddling terrestrial and maritime ecosystems triggered abrupt, synchronous, and multi-trophic ecological disruptions, including mortality, demographic shifts and altered species distributions. Tree die-off and coral bleaching occurred concurrently in response to the heat wave, and were accompanied by terrestrial plant mortality, seagrass and kelp loss, population crash of an endangered terrestrial bird species, plummeting breeding success in marine penguins, and outbreaks of terrestrial wood-boring insects. These multiple taxa and trophic-level impacts spanned >300,000 km2—comparable to the size of California—encompassing one terrestrial Global Biodiversity Hotspot and two marine World Heritage Areas. The subcontinental multi-taxa context documented here reveals that terrestrial and marine biotic responses to heat waves do not occur in isolation, implying that the extent of ecological vulnerability to projected increases in heat waves is underestimated.

A Dirty Dozen Ways to Die: Metrics and Modifiers of Mortality Driven by Drought and Warming for a Tree Species

Tree mortality events driven by drought and warmer temperature, often amplified by pests and pathogens, are emerging as one of the predominant climate change impacts on plants. Understanding and predicting widespread tree mortality events in the future is vital as they affect ecosystem goods and services provided by forests and woodlands, including carbon storage needed to help offset warming. Additionally, if extensive enough, tree die-off events can influence not only local climate but also climate and vegetation elsewhere via climate teleconnections. Consequently, recent efforts have focused on improving predictions of tree mortality. The most commercially important genera of trees is Pinus, and the most studied species for tree mortality is the pinon pine, Pinus edulis. Numerous metrics have been developed in association with predicting mortality thresholds or variations in mortality for this species. In this Mini Review we compiled metrics associated with drought and warming related mortality that were developed for P. edulis or for which P. edulis was a key example species used in a calculation or prediction. We grouped these metrics into four categories: (i) those related to simple climate variables, (ii) those related to physiological responses, (iii) those that require multi-step calculations or modeling using climate, ecohydrological, and/or ecophysiological data, and (iv) modifiers of rates or sensitivities of mortality. We identified the spatial-temporal scale of each of these metrics. The metrics to predict mortality include empirical ones which often have implicit linkages to expected mechanisms, and more mechanistic ones related to physiological drivers. The metrics for P. edulis have similarities with those available for other species of Pinus. Expected future mortality events will provide an opportunity to observationally and experimentally test and compare these metrics related to tree mortality for P. edulis via near-term ecological forecasting. The metrics for P. edulis may also be useful as potential analogs for other genera. Improving predictions of tree mortality for this species and others will be increasingly important as an aid to move towards anticipatory management.

Beyond greenness: Detecting temporal changes in photosynthetic capacity with hyperspectral reflectance data

Earths future carbon balance and regional carbon exchange dynamics are inextricably linked to plant photosynthesis. Spectral vegetation indices are widely used as proxies for vegetation greenness and to estimate state variables such as vegetation cover and leaf area index. However, the capacity of green leaves to take up carbon can change throughout the season. We quantify photosynthetic capacity as the maximum rate of RuBP carboxylation (Vcmax) and regeneration (Jmax). Vcmax and Jmax vary within-season due to interactions between ontogenetic processes and meteorological variables. Remote sensing-based estimation of Vcmax and Jmax using leaf reflectance spectra is promising, but temporal variation in relationships between these key determinants of photosynthetic capacity, leaf reflectance spectra, and the models that link these variables has not been evaluated. To address this issue, we studied hybrid poplar (Populus spp.) during a 7-week mid-summer period to quantify seasonally-dynamic relationships between Vcmax, Jmax, and leaf spectra. We compared in situ estimates of Vcmax and Jmax from gas exchange measurements to estimates of Vcmax and Jmax derived from partial least squares regression (PLSR) and fresh-leaf reflectance spectroscopy. PLSR models were robust despite dynamic temporal variation in Vcmax and Jmax throughout the study period. Within-population variation in plant stress modestly reduced PLSR model predictive capacity. Hyperspectral vegetation indices were well-correlated to Vcmax and Jmax, including the widely-used Normalized Difference Vegetation Index. Our results show that hyperspectral estimation of plant physiological traits using PLSR may be robust to temporal variation. Additionally, hyperspectral vegetation indices may be sufficient to detect temporal changes in photosynthetic capacity in contexts similar to those studied here. Overall, our results highlight the potential for hyperspectral remote sensing to estimate determinants of photosynthetic capacity during periods with dynamic temporal variations related to seasonality and plant stress, thereby improving estimates of plant productivity and characterization of the associated carbon budget.

Candidate halophytic grasses for addressing land degradation: Shoot responses of Sporobolus airoides and Paspalum vaginatum to weekly increasing NaCl concentration

ABSTRACT In many arid and semiarid regions worldwide, high levels of soil salinity is a key driver of land degradation, as well as a key impediment to re-establishing plant cover. Combating land degradation and erosion associated with soil salinity requires experimental determination of plant species that can grow in soils with high levels of salinity and can be used to re-establish plant cover. Herein, we evaluated the responses of untested candidate cultivars of two halophytic grass species to high soil salinity: alkali sacaton (Sporobolus airoides Torr.) and seashore paspalum (Paspalum vaginatum Swartz). We evaluated the growth responses of both species in a greenhouse under control (no-salt) and various levels of NaCl salinity (EC 8, 16, 24, 32, 40, and 48u2009dSm−1) using Hoagland solution in a hydroponics system in a randomized complete block design trial. At all salinity levels, sacaton grass had a greater shoot height, shorter root length, lower shoot fresh and dry weights, and poorer color and general quality compared to seashore paspalum. The shoot fresh and dry weights of both grasses were greatest at the low to medium levels of salinity, with the greatest response observed at EC 16u2009dSm−1. At the highest level, salinity significantly reduced shoot fresh and dry weights of both grasses. Because growth of both halophytic species exhibited high tolerance to salinity stress and were stimulated under low to medium levels of salinity, both species could be considered suitable candidates for re-establishing plant cover in drylands to combat desertification and land degradation associated with high levels of soil salinity.