How deregulation, drought and increasing fire impact Amazonian biodiversity.

Biodiversity contributes to the ecological and climatic stability of the Amazon Basin1,2, but is increasingly threatened by deforestation and fire3,4. Here we quantify these impacts over the past two decades using remote-sensing estimates of fire and deforestation and comprehensive range estimates of 11,514 plant species and 3,079 vertebrate species in the Amazon. Deforestation has led to large amounts of habitat loss, and fires further exacerbate this already substantial impact on Amazonian biodiversity. Since 2001, 103,079-189,755 km2 of Amazon rainforest has been impacted by fires, potentially impacting the ranges of 77.3-85.2% of species that are listed as threatened in this region5. The impacts of fire on the ranges of species in Amazonia could be as high as 64%, and greater impacts are typically associated with species that have restricted ranges. We find close associations between forest policy, fire-impacted forest area and their potential impacts on biodiversity. In Brazil, forest policies that were initiated in the mid-2000s corresponded to reduced rates of burning. However, relaxed enforcement of these policies in 2019 has seemingly begun to reverse this trend: approximately 4,253-10,343 km2 of forest has been impacted by fire, leading to some of the most severe potential impacts on biodiversity since 2009. These results highlight the critical role of policy enforcement in the preservation of biodiversity in the Amazon.

South American Dry Chaco rangelands: Positive effects of cattle trampling and transit on ecohydrological functioning.

Livestock production in drylands requires consideration of the ecological applications of ecohydrological redistribution of water. Intensive cattle trampling and the associated increase of surface runoff are common concerns for rangeland productivity and sustainability. Here, we highlight a regional livestock production system in which cattle trails and trampling surrounding an artificial impoundment are purposely managed to enhance redistribution and availability of water for cattle drinking. Based on literature synthesis and field measurements, we first describe cattle production systems and surface water redistribution in the Dry Chaco rangelands of South America, and then develop a conceptual framework to synthesize the ecohydrological impacts of livestock production on these ecosystems. Critical to this framework is the pioshere-a degraded overgrazed and overtrampled area where vegetation has difficulties growing, usually close to the water points. The Dry Chaco rangelands have three key distinctive characteristics associated with the flat sedimentary environment lacking fresh groundwater and the very extensive ranching conditions: (1) cattle drinking water is provided by artificial impoundments filled by runoff, (2) heavy trampling around the impoundment and its adjacent areas generates a piosphere that favors runoff toward the impoundment, and (3) the impoundment, piosphere, and extensive forage areas are hydrologically connected with a network of cattle trails. We propose an ecohydrological framework where cattle transit and trampling alter the natural water circulation of these ecosystems, affecting small fractions of the landscape through increased runoff (compaction in piosphere and trails), surface connectivity (convergence of trails to piosphere to impoundment), and ponding (compaction of the impoundment floor) that operate together making water harvesting and storage possible. These effects have likely generated a positive water feedback on the expansion of livestock in the region with a relatively low impact on forage production. We highlight the role of livestock transit as a geomorphological agent capable of reshaping the hydrology of flat sedimentary rangelands in ways that can be managed positively for sustainable ranching systems. We suggest that the Dry Chaco offers an alternative paradigm for rangelands in which cattle trampling may contribute to sustainable seminatural production systems with implications for other dry and flat rangelands of the world.

Climate-driven, but dynamic and complex? A reconciliation of competing hypotheses for species' distributions.

Estimates of the percentage of species "committed to extinction" by climate change range from 15% to 37%. The question is whether factors other than climate need to be included in models predicting species' range change. We created demographic range models that include climate vs. climate-plus-competition, evaluating their influence on the geographic distribution of Pinus edulis, a pine endemic to the semiarid southwestern U.S. Analyses of data on 23,426 trees in 1941 forest inventory plots support the inclusion of competition in range models. However, climate and competition together only partially explain this species' distribution. Instead, the evidence suggests that climate affects other range-limiting processes, including landscape-scale, spatial processes such as disturbances and antagonistic biotic interactions. Complex effects of climate on species distributions-through indirect effects, interactions, and feedbacks-are likely to cause sudden changes in abundance and distribution that are not predictable from a climate-only perspective.

Biological invasions and climate change amplify each other's effects on dryland degradation.

Climate models predict that, in the coming decades, many arid regions will experience increasingly hot conditions and will be affected more frequently by drought. These regions are also experiencing rapid vegetation change, notably invasion by exotic grasses. Invasive grasses spread rapidly into native desert ecosystems due, in particular, to interannual variability in precipitation and periodic fires. The resultant destruction of non-fire-adapted native shrub and grass communities and of the inherent soil resource heterogeneity can yield invader-dominated grasslands. Moreover, recurrent droughts are expected to cause widespread physiological stress and mortality of both invasive and native plants, as well as the loss of soil resources. However, the magnitude of these effects may differ between invasive and native grasses, especially under warmer conditions, rendering the trajectory of vegetated communities uncertain. Using the Biosphere 2 facility in the Sonoran Desert, we evaluated the viability of these hypothesized relationships by simulating combinations of drought and elevated temperature (+5°C) and assessing the ecophysiological and mortality responses of both a dominant invasive grass (Pennisetum ciliare or buffelgrass) and a dominant native grass (Heteropogan contortus or tanglehead). While both grasses survived protracted drought at ambient temperatures by inducing dormancy, drought under warmed conditions exceeded the tolerance limits of the native species, resulting in greater and more rapid mortality than exhibited by the invasive. Thus, two major drivers of global environmental change, biological invasion and climate change, can be expected to synergistically accelerate ecosystem degradation unless large-scale interventions are enacted.

Underappreciated plant vulnerabilities to heat waves.

With climate change, heat waves are becoming increasingly frequent, intense and broader in spatial extent. However, while the lethal effects of heat waves on humans are well documented, the impacts on flora are less well understood, perhaps except for crops. We summarize recent findings related to heat wave impacts including: sublethal and lethal effects at leaf and plant scales, secondary ecosystem effects, and more complex impacts such as increased heat wave frequency across all seasons, and interactions with other disturbances. We propose generalizable practical trials to quantify the critical bounding conditions of vulnerability to heat waves. Collectively, plant vulnerabilities to heat waves appear to be underappreciated and understudied, particularly with respect to understanding heat wave driven plant die-off and ecosystem tipping points.

Research frontiers for improving our understanding of drought-induced tree and forest mortality.

Accumulating evidence highlights increased mortality risks for trees during severe drought, particularly under warmer temperatures and increasing vapour pressure deficit (VPD). Resulting forest die-off events have severe consequences for ecosystem services, biophysical and biogeochemical land-atmosphere processes. Despite advances in monitoring, modelling and experimental studies of the causes and consequences of tree death from individual tree to ecosystem and global scale, a general mechanistic understanding and realistic predictions of drought mortality under future climate conditions are still lacking. We update a global tree mortality map and present a roadmap to a more holistic understanding of forest mortality across scales. We highlight priority research frontiers that promote: (1) new avenues for research on key tree ecophysiological responses to drought; (2) scaling from the tree/plot level to the ecosystem and region; (3) improvements of mortality risk predictions based on both empirical and mechanistic insights; and (4) a global monitoring network of forest mortality. In light of recent and anticipated large forest die-off events such a research agenda is timely and needed to achieve scientific understanding for realistic predictions of drought-induced tree mortality. The implementation of a sustainable network will require support by stakeholders and political authorities at the international level.

Rainfall intensity switches ecohydrological runoff/runon redistribution patterns in dryland vegetation patches.

Effectively managing net primary productivity in drylands for grazing and other uses depends on understanding how limited rainfall input is redistributed by runoff and runon among vegetation patches, particularly for patches that contrast between lesser and greater amounts of vegetation cover. Due in part to data limitations, ecohydrologists generally have focused on rainfall event size to characterize water redistribution processes. Here we use soil moisture data from a semiarid woodland to highlight how, when event size is controlled and runoff and interception are negligible at the stand scale, rainfall intensity drives the relationship between water redistribution and canopy and soil patch attributes. Horizontal water redistribution variability increased with rainfall intensity and differed between patches with contrasting vegetation cover. Sparsely vegetated patches gained relatively more water during lower intensity events, whereas densely vegetated ones gained relatively more water during higher intensity events. Consequently, range managers need to account for the distribution of rainfall event intensity, as well as event size, to assess the consequences of climate variability and change on net primary productivity. More generally, our results suggest that rainfall intensity needs to be considered in addition to event size to understand vegetation patch dynamics in drylands.

Precipitation thresholds and drought-induced tree die-off: insights from patterns of Pinus edulis mortality along an environmental stress gradient.

Recent regional tree die-off events appear to have been triggered by a combination of drought and heat - referred to as 'global-change-type drought'. To complement experiments focused on resolving mechanisms of drought-induced tree mortality, an evaluation of how patterns of tree die-off relate to highly spatially variable precipitation is needed. Here, we explore precipitation relationships with a die-off event of pinyon pine (Pinus edulis Engelm.) in southwestern North America during the 2002-2003 global-change-type drought. Pinyon die-off and its relationship with precipitation was quantified spatially along a precipitation gradient in north-central New Mexico with standard field plot measurements of die-off combined with canopy cover derived from normalized burn ratio (NBR) from Landsat imagery. Pinyon die-off patterns revealed threshold responses to precipitation (cumulative 2002-2003) and vapor pressure deficit (VPD), with little to no mortality (< 10%) above 600 mm and below warm season VPD of c. 1.7 kPa. [Correction added after online publication 17 June 2013; in the preceding sentence, the word 'below' has been inserted.] Our results refine how precipitation patterns within a region influence pinyon die-off, revealing a precipitation and VPD threshold for tree mortality and its uncertainty band where other factors probably come into play - a response type that influences stand demography and landscape heterogeneity and is of general interest, yet has not been documented.

Nonstructural leaf carbohydrate dynamics of Pinus edulis during drought-induced tree mortality reveal role for carbon metabolism in mortality mechanism.

Vegetation change is expected with global climate change, potentially altering ecosystem function and climate feedbacks. However, causes of plant mortality, which are central to vegetation change, are understudied, and physiological mechanisms remain unclear, particularly the roles of carbon metabolism and xylem function. We report analysis of foliar nonstructural carbohydrates (NSCs) and associated physiology from a previous experiment where earlier drought-induced mortality of Pinus edulis at elevated temperatures was associated with greater cumulative respiration. Here, we predicted faster NSC decline for warmed trees than for ambient-temperature trees. Foliar NSC in droughted trees declined by 30% through mortality and was lower than in watered controls. NSC decline resulted primarily from decreased sugar concentrations. Starch initially declined, and then increased above pre-drought concentrations before mortality. Although temperature did not affect NSC and sugar, starch concentrations ceased declining and increased earlier with higher temperatures. Reduced foliar NSC during lethal drought indicates a carbon metabolism role in mortality mechanism. Although carbohydrates were not completely exhausted at mortality, temperature differences in starch accumulation timing suggest that carbon metabolism changes are associated with time to death. Drought mortality appears to be related to temperature-dependent carbon dynamics concurrent with increasing hydraulic stress in P. edulis and potentially other similar species.

No carbon storage in growth-limited trees in a semi-arid woodland.

Plant survival depends on a balance between carbon supply and demand. When carbon supply becomes limited, plants buffer demand by using stored carbohydrates (sugar and starch). During drought, NSCs (non-structural carbohydrates) may accumulate if growth stops before photosynthesis. This expectation is pervasive, yet few studies have combined simultaneous measurements of drought, photosynthesis, growth, and carbon storage to test this. Using a field experiment with mature trees in a semi-arid woodland, we show that growth and photosynthesis slow in parallel as [Formula: see text] declines, preventing carbon storage in two species of conifer (J. monosperma and P. edulis). During experimental drought, growth and photosynthesis were frequently co-limited. Our results point to an alternative perspective on how plants use carbon that views growth and photosynthesis as independent processes both regulated by water availability.

Drought-induced forest decline: causes, scope and implications.

A large number of episodes of forest mortality associated with drought and heat stress have been detected worldwide in recent decades, suggesting that some of the world's forested ecosystems may be already responding to climate change. Here, we summarize a special session titled 'Drought-induced forest decline: causes, scope and implications' within the 12th European Ecological Federation Congress, held in Ávila (Spain) from 25 to 29 September 2011. The session focused on the interacting causes and impacts of die-off episodes at the community and ecosystem levels, and highlighted recent events of drought- and heat-related tree decline, advances in understanding mechanisms and in predicting mortality events, and diverse consequences of forest decline. Talks and subsequent discussion noted a potentially important role of carbon that may be interrelated with plant hydraulics in the multi-faceted process leading to drought-induced mortality; a substantial and yet understudied capacity of many forests to cope with extreme climatic events; and the difficulty of separating climate effects from other anthropogenic changes currently shaping forest dynamics in many regions of the Earth. The need for standard protocols and multi-level monitoring programmes to track the spatio-temporal scope of forest decline globally was emphasized as critical for addressing this emerging environmental issue.

Temperature sensitivity of drought-induced tree mortality portends increased regional die-off under global-change-type drought.

Large-scale biogeographical shifts in vegetation are predicted in response to the altered precipitation and temperature regimes associated with global climate change. Vegetation shifts have profound ecological impacts and are an important climate-ecosystem feedback through their alteration of carbon, water, and energy exchanges of the land surface. Of particular concern is the potential for warmer temperatures to compound the effects of increasingly severe droughts by triggering widespread vegetation shifts via woody plant mortality. The sensitivity of tree mortality to temperature is dependent on which of 2 non-mutually-exclusive mechanisms predominates--temperature-sensitive carbon starvation in response to a period of protracted water stress or temperature-insensitive sudden hydraulic failure under extreme water stress (cavitation). Here we show that experimentally induced warmer temperatures (approximately 4 degrees C) shortened the time to drought-induced mortality in Pinus edulis (piñon shortened pine) trees by nearly a third, with temperature-dependent differences in cumulative respiration costs implicating carbon starvation as the primary mechanism of mortality. Extrapolating this temperature effect to the historic frequency of water deficit in the southwestern United States predicts a 5-fold increase in the frequency of regional-scale tree die-off events for this species due to temperature alone. Projected increases in drought frequency due to changes in precipitation and increases in stress from biotic agents (e.g., bark beetles) would further exacerbate mortality. Our results demonstrate the mechanism by which warmer temperatures have exacerbated recent regional die-off events and background mortality rates. Because of pervasive projected increases in temperature, our results portend widespread increases in the extent and frequency of vegetation die-off.

Global field observations of tree die-off reveal hotter-drought fingerprint for Earth's forests.

Earth's forests face grave challenges in the Anthropocene, including hotter droughts increasingly associated with widespread forest die-off events. But despite the vital importance of forests to global ecosystem services, their fates in a warming world remain highly uncertain. Lacking is quantitative determination of commonality in climate anomalies associated with pulses of tree mortality-from published, field-documented mortality events-required for understanding the role of extreme climate events in overall global tree die-off patterns. Here we established a geo-referenced global database documenting climate-induced mortality events spanning all tree-supporting biomes and continents, from 154 peer-reviewed studies since 1970. Our analysis quantifies a global "hotter-drought fingerprint" from these tree-mortality sites-effectively a hotter and drier climate signal for tree mortality-across 675 locations encompassing 1,303 plots. Frequency of these observed mortality-year climate conditions strongly increases nonlinearly under projected warming. Our database also provides initial footing for further community-developed, quantitative, ground-based monitoring of global tree mortality.

Interactive effects of grazing and burning on wind- and water-driven sediment fluxes: rangeland management implications.

Rangelands are globally extensive, provide fundamental ecosystem services, and are tightly coupled human-ecological systems. Rangeland sustainability depends largely on the implementation and utilization of various grazing and burning practices optimized to protect against soil erosion and transport. In many cases, however, land management practices lead to increased soil erosion and sediment fluxes for reasons that are poorly understood. Because few studies have directly measured both wind and water erosion and transport, an assessment of how they may differentially respond to grazing and burning practices is lacking. Here, we report simultaneous, co-located estimates of wind- and water-driven sediment transport in a semiarid grassland in Arizona, USA, over three years for four land management treatments: control, grazed, burned, and burned + grazed. For all treatments and most years, annual rates of wind-driven sediment transport exceeded that of water due to a combination of ongoing small but nontrivial wind events and larger, less frequent, wind events that generally preceded the monsoon season. Sediment fluxes by both wind and water differed consistently by treatment: burned + grazed > burned >> grazed > or = control, with effects immediately apparent after burning but delayed after grazing until the following growing season. Notably, the wind:water sediment transport ratio decreased following burning but increased following grazing. Our results show how rangeland practices disproportionally alter sediment fluxes driven by wind and water, differences that could potentially help explain divergence between rangeland sustainability and degradation.

When ecosystem services crash: preparing for big, fast, patchy climate change.

Assessments of adaptation options generally focus on incremental, homogeneous ecosystem responses to climate even though climate change impacts can be big, fast, and patchy across a region. Regional drought-induced tree die-off in semiarid woodlands highlights how an ecosystem crash fundamentally alters most ecosystem services and poses management challenges. Building on previous research showing how choice of location is linked to adaptive capacity and vulnerability, we developed a framework showing how the options for retaining desired ecosystem services in the face of sudden crashes depend on how portable the service is and whether the stakeholder is flexible with regard to the location where they receive their services. Stakeholders using portable services, or stakeholders who can move to other locations to obtain services, may be more resilient to ecosystem crashes. Our framework suggests that entering into cooperative networks with regionally distributed stakeholders is key to building resilience to big, fast, patchy crashes.

Radionuclide resuspension across ecosystems and environmental disturbances.

Exposure assessment from radionuclides and other soil-bound contaminants often requires quantifying the amount of contaminant resuspended in the air. Rates and controlling factors of radionuclide resuspension and wind erosion of soil are clearly related but have largely been studied separately. Here, we review both and then integrate wind erosion measurements with the radiological resuspension paradigm to provide better estimates of resuspension factors across a broad range of ecosystems and environmental conditions. Radionuclide resuspension by wind was initially investigated during the era of aboveground nuclear weapons testing. Predictive dose models were developed from empirically-derived ratios of air and soil concentrations, otherwise called the resuspension factor. Resuspension factors were shown to generally predict radionuclide concentrations in air, but they were site-specific and largely derived from the arid and semi-arid environments surrounding nuclear weapons testing locations. In contrast, wind erosion studies from the agricultural and environmental sciences have produced more mechanistic models and a relatively robust data set of wind erosion rates and model parameters across a range of ecosystems. We sequentially show the mathematics linking measured sediment flux from wind erosion rate measurements to resuspension factors using the concept of transport capacity and its relationship to the deposition velocity. We also describe the conceptual framework describing how resuspension factors change through time and the mathematical models describing this decrease. We then show how vertical mass flux measurements across ecosystems were categorized and used to calculate ecosystem-based resuspension factors. These calculations allow generalized estimation of radionuclide resuspension factors across ecosystem types as a function of disturbance and as input for dose calculations.

Vegetation responses to extreme hydrological events: sequence matters.

Extreme hydrological events such as flood and drought drive vegetation dynamics and are projected to increase in frequency in association with climate change, which could result in sequences of extreme events. However, experimental studies of vegetation responses to climate have largely focused on responses to a trend in climate or to a single extreme event but have largely overlooked the potential for complex responses to specific sequences of extreme events. Here we document, on the basis of an experiment with seedlings of three types of subtropical wetland tree species, that mortality can be amplified and growth can even be stimulated, depending on event sequence. Our findings indicate that the impacts of multiple extreme events cannot be modeled by simply summing the projected effects of individual extreme events but, rather, that models should take into account event sequences.

Mechanisms of plant survival and mortality during drought: why do some plants survive while others succumb to drought?

Severe droughts have been associated with regional-scale forest mortality worldwide. Climate change is expected to exacerbate regional mortality events; however, prediction remains difficult because the physiological mechanisms underlying drought survival and mortality are poorly understood. We developed a hydraulically based theory considering carbon balance and insect resistance that allowed development and examination of hypotheses regarding survival and mortality. Multiple mechanisms may cause mortality during drought. A common mechanism for plants with isohydric regulation of water status results from avoidance of drought-induced hydraulic failure via stomatal closure, resulting in carbon starvation and a cascade of downstream effects such as reduced resistance to biotic agents. Mortality by hydraulic failure per se may occur for isohydric seedlings or trees near their maximum height. Although anisohydric plants are relatively drought-tolerant, they are predisposed to hydraulic failure because they operate with narrower hydraulic safety margins during drought. Elevated temperatures should exacerbate carbon starvation and hydraulic failure. Biotic agents may amplify and be amplified by drought-induced plant stress. Wet multidecadal climate oscillations may increase plant susceptibility to drought-induced mortality by stimulating shifts in hydraulic architecture, effectively predisposing plants to water stress. Climate warming and increased frequency of extreme events will probably cause increased regional mortality episodes. Isohydric and anisohydric water potential regulation may partition species between survival and mortality, and, as such, incorporating this hydraulic framework may be effective for modeling plant survival and mortality under future climate conditions.

Phenology of mixed woody-herbaceous ecosystems following extreme events: net and differential responses.

Ecosystem responses to key climate drivers are reflected in phenological dynamics such as the timing and degree of "green-up" that integrate responses over spatial scales from individual plants to ecosystems. This integration is clearest in ecosystems dominated by a single species or life form, such as seasonally dynamic grasslands or more temporally constant evergreen forests. Yet many ecosystems have substantial contribution of cover from both herbaceous and woody evergreen plants. Responses of mixed woody-herbaceous ecosystems to climate are of increasing concern due to their extensive nature, the potential for such systems to yield more complex responses than those dominated by a single life form, and projections that extreme climate and weather events will increase in frequency and intensity with global warming. We present responses of a mixed woody-herbaceous ecosystem type to an extreme event: regional-scale piñon pine mortality following an extended drought and the subsequent herbaceous green-up following the first wet period after the drought. This example highlights how reductions in greenness of the slower, more stable evergreen woody component can rapidly be offset by increases associated with resources made available to the relatively more responsive herbaceous component. We hypothesize that such two-phase phenological responses to extreme events are characteristic of many mixed woody-herbaceous ecosystems.

Foliar absorption of intercepted rainfall improves woody plant water status most during drought.

A large proportion of rainfall in dryland ecosystems is intercepted by plant foliage and is generally assumed to evaporate to the atmosphere or drip onto the soil surface without being absorbed. We demonstrate foliar absorption of intercepted rainfall in a widely distributed, continental dryland, woody-plant genus: Juniperus. We observed substantial improvement in plant water status, exceeding 1.0 MPa water potential for drought-stressed plants, following precipitation on an experimental plot that excluded soil water infiltration. Experiments that wetted shoots with unlabeled and with isotopically labeled water confirmed that water potential responded substantially to foliar wetting, that these responses were not attributable to re-equilibration with other portions of the xylem, and that magnitude of response increased with water stress. Foliar absorption is not included in most ecological, hydrological, and atmospheric models; has implications for interpreting plant isotopic signatures; and not only supplements water acquisition associated with increases in soil moisture that follow large or repeated precipitation events, but also enables plants to bypass soil water uptake and benefit from the majority of precipitation events, which wet foliage but do not increase soil moisture substantially. Foliar absorption of intercepted water could be more important than previously appreciated, especially during drought when water stress is greatest.