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Patterns and causes of observed piñon pine mortality in the southwestern United States

Recently, widespread piñon pine die-off occurred in the southwestern United States. Here we synthesize observational studies of this event and compare findings to expected relationships with biotic and abiotic factors. Agreement exists on the occurrence of drought, presence of bark beetles and increased mortality of larger trees. However, studies disagree about the influences of stem density, elevation and other factors, perhaps related to study design, location and impact of extreme drought. Detailed information about bark beetles is seldom reported and their role is poorly understood. Our analysis reveals substantial limits to our knowledge regarding the processes that produce mortality patterns across space and time, indicating a poor ability to forecast mortality in response to expected increases in future droughts.

New Phytologist

Moisture rivals temperature in limiting photosynthesis by trees establishing beyond their cold-edge range limit under ambient and warmed conditions

Climate change is altering plant species distributions globally, and warming is expected to promote uphill shifts in mountain trees. However, at many cold-edge range limits, such as alpine treelines in the western United States, tree establishment may be colimited by low temperature and low moisture, making recruitment patterns with warming difficult to predict. We measured response functions linking carbon (C) assimilation and temperature- and moisture-related microclimatic factors for limber pine ( Pinus flexilis ) seedlings growing in a heating × watering experiment within and above the alpine treeline. We then extrapolated these response functions using observed microclimate conditions to estimate the net effects of warming and associated soil drying on C assimilation across an entire growing season. Moisture and temperature limitations were each estimated to reduce potential growing season C gain from a theoretical upper limit by 15–30% ( c . 50% combined). Warming above current treeline conditions provided relatively little benefit to modeled net assimilation, whereas assimilation was sensitive to either wetter or drier conditions. Summer precipitation may be at least as important as temperature in constraining C gain by establishing subalpine trees at and above current alpine treelines as seasonally dry subalpine and alpine ecosystems continue to warm.

Colorado

Tree mortality from drought, insects, and their interactions in a changing climate

Climate change is expected to drive increased tree mortality through drought, heat stress, and insect attacks, with manifold impacts on forest ecosystems. Yet, climate-induced tree mortality and biotic disturbance agents are largely absent from process-based ecosystem models. Using data sets from the western USA and associated studies, we present a framework for determining the relative contribution of drought stress, insect attack, and their interactions, which is critical for modeling mortality in future climates. We outline a simple approach that identifies the mechanisms associated with two guilds of insects – bark beetles and defoliators – which are responsible for substantial tree mortality. We then discuss cross-biome patterns of insect-driven tree mortality and draw upon available evidence contrasting the prevalence of insect outbreaks in temperate and tropical regions. We conclude with an overview of tools and promising avenues to address major challenges. Ultimately, a multitrophic approach that captures tree physiology, insect populations, and tree–insect interactions will better inform projections of forest ecosystem responses to climate change.

New Phytologist

Incorporating phosphorus cycling into global modeling efforts: a worthwhile, tractable endeavor

Myriad field, laboratory, and modeling studies show that nutrient availability plays a fundamental role in regulating CO 2 exchange between the Earth's biosphere and atmosphere, and in determining how carbon pools and fluxes respond to climatic change. Accordingly, global models that incorporate coupled climate–carbon cycle feedbacks made a significant advance with the introduction of a prognostic nitrogen cycle. Here we propose that incorporating phosphorus cycling represents an important next step in coupled climate–carbon cycling model development, particularly for lowland tropical forests where phosphorus availability is often presumed to limit primary production. We highlight challenges to including phosphorus in modeling efforts and provide suggestions for how to move forward.

New Phytologist

Towards understanding resprouting at the global scale

Understanding and predicting plant response to disturbance is of paramount importance in our changing world. Resprouting ability is often considered a simple qualitative trait and used in many ecological studies. Our aim is to show some of the complexities of resprouting while highlighting cautions that need be taken in using resprouting ability to predict vegetation responses across disturbance types and biomes. There are marked differences in resprouting depending on the disturbance type, and fire is often the most severe disturbance because it includes both defoliation and lethal temperatures. In the Mediterranean biome, there are differences in functional strategies to cope with water deficit between resprouters (dehydration avoiders) and nonresprouters (dehydration tolerators); however, there is little research to unambiguously extrapolate these results to other biomes. Furthermore, predictions of vegetation responses to changes in disturbance regimes require consideration not only of resprouting, but also other relevant traits (e.g. seeding, bark thickness) and the different correlations among traits observed in different biomes; models lacking these details would behave poorly at the global scale. Overall, the lessons learned from a given disturbance regime and biome (e.g. crown-fire Mediterranean ecosystems) can guide research in other ecosystems but should not be extrapolated at the global scale.

New Phytologist

Terrestrial nitrogen cycling in Earth system models revisited

Understanding the degree to which nitrogen (N) availability limits land carbon (C) uptake under global environmental change represents an unresolved challenge. First-generation ‘C-only’vegetation models, lacking explicit representations of N cycling,projected a substantial and increasing land C sink under rising atmospheric CO 2 concentrations. This prediction was questioned for not taking into account the potentially limiting effect of N availability, which is necessary for plant growth (Hungate et al.,2003). More recent global models include coupled C and N cycles in land ecosystems (C–N models) and are widely assumed to be more realistic. However, inclusion of more processes has not consistently improved their performance in capturing observed responses of the global C cycle (e.g. Wenzel et al., 2014). With the advent of a new generation of global models, including coupled C, N, and phosphorus (P) cycling, model complexity is sure to increase; but model reliability may not, unless greater attention is paid to the correspondence of model process representations ande mpirical evidence. It was in this context that the ‘Nitrogen Cycle Workshop’ at Dartington Hall, Devon, UK was held on 1–5 February 2016. Organized by I. Colin Prentice and Benjamin D. Stocker (Imperial College London, UK), the workshop was funded by the European Research Council,project ‘Earth system Model Bias Reduction and assessing Abrupt Climate change’ (EMBRACE). We gathered empirical ecologists and ecosystem modellers to identify key uncertainties in terrestrial C–N cycling, and to discuss processes that are missing or poorly represented in current models.

New Phytologist

Climate drives shifts in grass reproductive phenology across the western USA

The capacity of grass species to alter their reproductive timing across space and through time can indicate their ability to cope with environmental variability and help predict their future performance under climate change. We determined the long-term (1895–2013) relationship between flowering times of grass species and climate in space and time using herbarium records across ecoregions of the western USA. There was widespread concordance of C 3 grasses accelerating flowering time and general delays for C 4 grasses with increasing mean annual temperature, with the largest changes for annuals and individuals occurring in more northerly, wetter ecoregions. Flowering time was delayed for most grass species with increasing mean annual precipitation across space, while phenology–precipitation relationships through time were more mixed. Our results suggest that the phenology of most grass species has the capacity to respond to increases in temperature and altered precipitation expected with climate change, but weak relationships for some species in time suggest that climate tracking via migration or adaptation may be required. Divergence in phenological responses among grass functional types, species, and ecoregions suggests that climate change will have unequal effects across the western USA.

New Phytologist

Biocrust ecology: Unifying micro- and macro-scales to confront global change

Biological soil crusts (biocrusts) are communities of microbes, lichens and bryophytes living at the soil surface in drylands (Fig. 1; Belnap et al ., 2016). Biocrusts occur on all continents and can comprise a majority of cover in some systems (Belnap et al ., 2016). While species diversity and distributions have long been a research focus, interest in controls on community composition and cover has expanded as biocrusts are increasingly recognized for their roles in ecosystem functioning (Deane-Coe and Stanton, 2017). For example, biocrust organisms can stabilize soils (Belnap et al ., 2016; Faist et al ., 2017), fix atmospheric carbon (C) (Sancho et al ., 2016), and serve as the foremost source of ‘new’ soil nitrogen (N) in drylands, via N 2 fixation (Barger et al ., 2016) These contributions to gross primary production and soil fertility could be quite large, as high-end estimates suggest biocrusts and similar communities of bryophytes and lichens might account for 10% of terrestrial C- and 50% of N-fixation globally (Elbert et al ., 2012). Yet verifying these and other biocrust roles in ecosystem functioning is complicated by limited knowledge of biocrust cover and composition across the vast dryland biome (Ferrenberg et al ., 2017). It was against this backdrop that ‘Biocrust3: the 3 rd International workshop on biological soil crusts’ was held in Moab, UT, USA, on 26-30 September 2016. The workshop brought together over 50 scientists from 21 countries and six continents, and included numerous biocrust science pioneers (Fig. 2). The meeting was notable for its cross-scale focus, discussion of novel molecular and imaging techniques, and sessions on mapping and restoring biocrusts in a changing world. Here, we synthesize a central theme that emerged from Biocrust3, namely the potential for combining cutting edge tools with studies focused on organismal traits, ecosystem functions, and global change biology to advance the frontier of biocrust ecology.

New Phytologist

Monitoring global tree mortality patterns and trends. Report from the VW symposium ‘Crossing scales and disciplines to identify global trends of tree mortality as indicators of forest health’

From the 21stto the 23rdJune 2017, the Herrenhausen castle inHannover/Germany hosted a diverse and large crowd with morethan 70 tree physiologists, forest ecologists, forest inventoryexperts, remote-sensing scientists, and vegetation modele rs. Par-ticipants from six continent s and from more than 20 countriesgathered to discuss how to improve the scientific determination ofglobal-scale patterns, drivers, and trends of a threatening phe-nomenon: the apparent emergence of recent widespread treemortality events in diverse forests around the world.Continuin g the theme of a workshop held at the Max-PlanckInstitute for Biogeochemistry in Jena (Germany) in 2014 (Hart-mann et al., 2015), the Hanover meeting intended to developapproaches, tools and collaborative actions to accelerate progress inaddressing regional patterns and trends of tree mortality (Williamset al., 2013). Over the last decade climate change related treemortality events have been increasingly reported around the globe(van Mantgem et al., 2009; Carnicer et al., 2011; Peng et al., 2011;Brienen et al., 2015), but to what degree this is a global trend,amplifying under increasing climate change, remains uncertain.

New Phytologist

Drivers and mechanisms of tree mortality in moist tropical forests

Tree mortality rates appear to be increasing in moist tropical forests (MTFs) with significant carbon cycle consequences. Here, we review the state of knowledge regarding MTF tree mortality, create a conceptual framework with testable hypotheses regarding the drivers, mechanisms and interactions that may underlie increasing MTF mortality rates, and identify the next steps for improved understanding and reduced prediction. Increasing mortality rates are associated with rising temperature and vapor pressure deficit, liana abundance, drought, wind events, fire and, possibly, CO 2 fertilization‐induced increases in stand thinning or acceleration of trees reaching larger, more vulnerable heights. The majority of these mortality drivers may kill trees in part through carbon starvation and hydraulic failure. The relative importance of each driver is unknown. High species diversity may buffer MTFs against large‐scale mortality events, but recent and expected trends in mortality drivers give reason for concern regarding increasing mortality within MTFs. Models of tropical tree mortality are advancing the representation of hydraulics, carbon and demography, but require more empirical knowledge regarding the most common drivers and their subsequent mechanisms. We outline critical datasets and model developments required to test hypotheses regarding the underlying causes of increasing MTF mortality rates, and improve prediction of future mortality under climate change.

New Phytologist

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.

New Phytologist

Ecosystem thresholds, tipping points, and critical transitions

Abrupt shifts in ecosystems are cause for concern and will likelyintensify under global change (Scheffer et al., 2001). The terms‘thresho lds’, ‘tipping points’, and ‘critical transitions’ have beenused interchangeably to refer to sudden changes in the integrityor state of an ecosystem caused by environmental drivers(Holling, 1973; May, 1977). Threshold-based concepts havesignific antly aided our capacity to predict the controls overecosystem structure and functioning (Schwinning et al., 2004;Peters et al., 2007) and have become a framework to guide themanagement of natural resources (Glick et al., 2010; Allen et al.,2011). However, our unders tanding of how biotic and abioticdrivers interact to regulate ecosystem responses and of ways toforecast th e impending responses remain limited. Terrestrialecosystems, in particular, are already responding to globalchange in ways that are both transformati onal and difficult topredict due to strong heterogeneity across temporal and spatialscales (Pe~nuelas & Filella, 2001; McDowell et al., 2011;Munson, 2013; Reed et al., 2016). Comparing approaches formeasuring ecosystem performance in response to changingenvironme ntal conditions and for detecting stress and thresholdresponses can improve tradition al tests of resilience and provideearly warning signs of ecosystem transitions. Similarly, com-paring responses across ecosystems can offer insight into themechanisms that underlie variation in threshold responses.

New Phytologist

Science at the frontier: Multimethod research to evaluate ecosystem change across multiple scales

Changes in the Earth system occur across the full spectrum of spatial and temporal scales, yet our research approaches to understanding and predicting those changes are typically restricted to a pre-defined window of space and time. For this reason, there is substantial power in integrating different approaches, particularly for research associated with the multifaceted nature of ecosystem responses to global change. Within a given research approach – for example, remote sensing, field experimentation, modeling – science promotes the continued advancement of tools and techniques. As technical advancements continue at an unprecedented rate, new opportunities for integrated, multi-approach research emerge, which could more effectively capture the mechanisms and patterns that drive ecosystem structure and function. A capacity to move beyond comparison and into the realm of integration shows promise for promoting significant advances in Earth system science, as evidenced by the Organized Session Science at the Frontier: Using Multimethod Research to Create New Knowledge and Assess Tools Across Spatial and Temporal Scales on December 12, 2017 at the Fall meeting of the American Geophysical Union in New Orleans, Louisiana.

New Phytologist

Projected warming disrupts the synchrony of riparian seed dispersal and snowmelt streamflow

• Globally, spring phenology and abiotic processes are shifting earlier with warming. Differences in the magnitudes of these shifts may decouple the timing of plant resource requirements from resource availability. In riparian forests across the northern hemisphere, warming could decouple seed dispersal from snowmelt peak streamflow, thus reducing moisture and safe-sites for dominant tree recruitment. • We combined field observations with climate, hydrology, and phenology models to simulate future change in synchrony of seed dispersal and snowmelt peaks in the upper South Platte River Basin, Colorado, for three Salicaceae species that dominate western USA riparian forests. • Chilling requirements for overcoming winter endodormancy were strongest in Salix exigua, moderately supported for Populus deltoides, and indiscernible in Salix amygdaloides. Ensemble mean projected warming of 3.5ºC shifted snowmelt peaks 10-19 d earlier relative to S. exigua and P. deltoides dispersal, because decreased winter chilling combined with increased spring forcing limited change in their dispersal phenology. In contrast, warming shifted both snowmelt peaks and S. amygdaloides dispersal 21 d earlier, maintaining their synchrony. • Decoupling of snowmelt from seed dispersal for Salicaceae with strong chilling requirements is likely to reduce resources critical for recruitment of these foundational riparian forests, although the magnitude of future decoupling remains uncertain.

Colorado

Biogeography and phylogeny of masting: Do global patterns fit functional hypotheses?

1) Interannual variability of seed crops (CVp) has profound consequences for plant populations and food webs, where high CVp is termed ‘masting’. Here we ask: is global variation in CVp better predicted by plant or habitat differences consistent with adaptive economies of scale, in which flower and seed benefits increase disproportionately during mast years; or to passive mechanisms, in which seed production responds to variation in resource availability associated with climate variability? 2) To address this question, we compiled a dataset for phylogenetic comparative analysis of long-term fruit/seed production for plants comprising 920 time-series spanning 311 plant species. 3) Factors associated with both adaptive benefits of CVp (wind pollination and seed dispersal) and climatic variability (variability of summer precipitation) were among the best predictors of global variation in CVp. We observed a hump-shaped relationship between CVp and latitude and intermediate phylogenetic and geographic signals in CVp. 4) CVp is patterned non-randomly across the globe and over the plant tree of life, where high CVp is associated with species benefiting from economies of scale of seed or flower production and with species that experience variable rainfall over summer months when seeds usually mature.

New Phytologist

Mortality predispositions of conifers across western USA

Conifer mortality rates are increasing in western North America, but the physiological mechanisms underlying this trend are not well understood. We examined tree‐ring‐based radial growth along with stable carbon (C) and oxygen (O) isotope composition ( δ 13 C and δ 18 O, respectively) of dying and surviving conifers at eight old‐growth forest sites across a strong moisture gradient in the western USA to retrospectively investigate mortality predispositions. Compared with surviving trees, lower growth of dying trees was detected at least one decade before mortality at seven of the eight sites. Intrinsic water‐use efficiency increased over time in both dying and surviving trees, with a weaker increase in dying trees at five of the eight sites. C starvation was a strong correlate of conifer mortality based on a conceptual model incorporating growth, δ 13 C, and δ 18 O. However, this approach does not capture processes that occur in the final months of survival. Ultimately, C starvation may lead to increased mortality vulnerability, but hydraulic failure or biotic attack may dominate the process during the end stages of mortality in these conifers.

New Phytologist

Integrating the evidence for a terrestrial carbon sink caused by increasing atmospheric CO2

Atmospheric carbon dioxide concentration ([CO 2 ]) is increasing, which increases leaf‐scale photosynthesis and intrinsic water‐use efficiency. These direct responses have the potential to increase plant growth, vegetation biomass, and soil organic matter; transferring carbon from the atmosphere into terrestrial ecosystems (a carbon sink). A substantial global terrestrial carbon sink would slow the rate of [CO 2 ] increase and thus climate change. However, ecosystem CO 2 responses are complex or confounded by concurrent changes in multiple agents of global change and evidence for a [CO 2 ]‐driven terrestrial carbon sink can appear contradictory. Here we synthesize theory and broad, multidisciplinary evidence for the effects of increasing [CO 2 ] (iCO 2 ) on the global terrestrial carbon sink. Evidence suggests a substantial increase in global photosynthesis since pre‐industrial times. Established theory, supported by experiments, indicates that iCO 2 is likely responsible for about half of the increase. Global carbon budgeting, atmospheric data, and forest inventories indicate a historical carbon sink, and these apparent iCO 2 responses are high in comparison to experiments and predictions from theory. Plant mortality and soil carbon iCO 2 responses are highly uncertain. In conclusion, a range of evidence supports a positive terrestrial carbon sink in response to iCO 2 , albeit with uncertain magnitude and strong suggestion of a role for additional agents of global change.

New Phytologist

Biotic vs abiotic controls on temporal sensitivity of primary production to precipitation across North American drylands

Dryland net primary productivity (NPP) is sensitive to temporal variation in precipitation (PPT), but the magnitude of this ‘temporal sensitivity’ varies spatially. Hypotheses for spatial variation in temporal sensitivity have often emphasized abiotic factors, such as moisture limitation, while overlooking biotic factors, such as vegetation structure. We tested these hypotheses using spatiotemporal models fit to remote-sensing data sets to assess how vegetation structure and climate influence temporal sensitivity across five dryland ecoregions of the western USA. Temporal sensitivity was higher in locations and ecoregions dominated by herbaceous vegetation. By contrast, much less spatial variation in temporal sensitivity was explained by mean annual PPT. In fact, ecoregion-specific models showed inconsistent associations of sensitivity and PPT; whereas sensitivity decreased with increasing mean annual PPT in most ecoregions, it increased with mean annual PPT in the most arid ecoregion, the hot deserts. The strong, positive influence of herbaceous vegetation on temporal sensitivity indicates that herbaceous-dominated drylands will be particularly sensitive to future increases in precipitation variability and that dramatic changes in cover type caused by invasions or shrub encroachment will lead to changes in dryland NPP dynamics, perhaps independent of changes in precipitation.

New Phytologist