A B S T R A C T Greenhouse gas emissions have significantly altered global climate, and will continue to do so in the future. Increases in the frequency, duration, and/or severity of drought and heat stress associated with climate change could fundamentally alter the composition, structure, and biogeography of forests in many regions. Of particular concern are potential increases in tree mortality associated with climateinduced physiological stress and interactions with other climate-mediated processes such as insect outbreaks and wildfire. Despite this risk, existing projections of tree mortality are based on models that lack functionally realistic mortality mechanisms, and there has been no attempt to track observations of climate-driven tree mortality globally. Here we present the first global assessment of recent tree mortality attributed to drought and heat stress. Although episodic mortality occurs in the absence of climate change, studies compiled here suggest that at least some of the world's forested ecosystems already may be responding to climate change and raise concern that forests may become increasingly vulnerable to higher background tree mortality rates and die-off in response to future warming and drought, even in environments that are not normally considered water-limited. This further suggests risks to ecosystem services, including the loss of sequestered forest carbon and associated atmospheric feedbacks. Our review also identifies key information gaps and scientific uncertainties that currently hinder our ability to predict tree mortality in response to climate change and emphasizes the need for a globally coordinated observation system. Overall, our review reveals the potential for amplified tree mortality due to drought and heat in forests worldwide.Published by Elsevier B.V.

Plant traits-the morphological, anatomical, physiological, biochemical and phenological characteristics of plants-determine how plants respond to environmental factors, affect other trophic levels, and influence ecosystem properties and their benefits and detriments to people. Plant trait data thus represent the basis for a vast area of research spanning from evolutionary biology, community and functional ecology, to biodiversity conservation, ecosystem and landscape management, restoration, biogeography and earth system modelling. Since its foundation in 2007, the TRY database of plant traits has grown continuously. It now provides unprecedented data coverage under an open access data policy and is the main plant trait database used by the research community worldwide. Increasingly, the TRY database also supports new frontiers of trait-based plant research, including the identification of data gaps and the subsequent mobilization or measurement of new data. To support this development, in this article we evaluate the extent of the trait data compiled in TRY and analyse emerging patterns of data coverage and representativeness. Best species coverage is achieved for categorical traits-almost complete coverage for 'plant growth form'. However, most traits relevant for ecology and vegetation modelling are characterized by continuous intraspecific variation and trait-environmental relationships. These traits have to be measured on individual plants in their respective environment. Despite unprecedented data coverage, we observe a humbling lack of completeness and representativeness of these continuous traits in many aspects.We, therefore, conclude that reducing data gaps and biases in the TRY database remains a key challenge and requires a coordinated approach to data mobilization and trait measurements. This can only be achieved in collaboration with other initiatives. Geosphere-Biosphere Program (IGBP) and DIVERSITAS, the TRY database (TRY-not an acronym, rather a statement of sentiment; https ://www.try-db.org; Kattge et al., 2011) was proposed with the explicit assignment to improve the availability and accessibility of plant trait data for ecology and earth system sciences. The Max Planck Institute for Biogeochemistry (MPI-BGC) offered to host the database and the different groups joined forces for this community-driven program. Two factors were key to the success of TRY: the support and trust of leaders in the field of functional plant ecology submitting large databases and the long-term funding by the Max Planck Society, the MPI-BGC and the German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, which has enabled the continuous development of the TRY database.

Tree mortality is a key factor influencing forest functions and dynamics, but our understanding of the mechanisms leading to mortality and the associated changes in tree growth rates are still limited. We compiled a new pan-continental tree-ring width database from sites where both dead and living trees were sampled (2970 dead and 4224 living trees from 190 sites, including 36 species), and compared early and recent growth rates between trees that died and those that survived a given mortality event. We observed a decrease in radial growth before death in ca. 84% of the mortality events. The extent and duration of these reductions were highly variable (1-100 years in 96% of events) due to the complex interactions among study species and the source(s) of mortality. Strong and long-lasting declines were found for gymnosperms, shade- and drought-tolerant species, and trees that died from competition. Angiosperms and trees that died due to biotic attacks (especially bark-beetles) typically showed relatively small and short-term growth reductions. Our analysis did not highlight any universal trade-off between early growth and tree longevity within a species, although this result may also reflect high variability in sampling design among sites. The intersite and interspecific variability in growth patterns before mortality provides valuable information on the nature of the mortality process, which is consistent with our understanding of the physiological mechanisms leading to mortality. Abrupt changes in growth immediately before death can be associated with generalized hydraulic failure and/or bark-beetle attack, while long-term decrease in growth may be associated with a gradual decline in hydraulic performance coupled with depletion in carbon reserves. Our results imply that growth-based mortality algorithms may be a powerful tool for predicting gymnosperm mortality induced by chronic stress, but not necessarily so for angiosperms and in case of intense drought or bark-beetle outbreaks.

Drought events are increasing globally, and reports of consequent forest mortality are widespread. However, due to a lack of a quantitative global synthesis, it is still not clear whether droughtinduced mortality rates differ among global biomes and whether functional traits influence the risk of drought-induced mortality. To address these uncertainties, we performed a global metaanalysis of 58 studies of drought-induced forest mortality. Mortality rates were modelled as a function of drought, temperature, biomes, phylogenetic and functional groups and functional traits. We identified a consistent global-scale response, where mortality increased with drought severity [log mortality (trees trees À1 year À1 ) increased 0.46 (95% CI = 0.2-0.7) with one SPEI unit drought intensity]. We found no significant differences in the magnitude of the response depending on forest biomes or between angiosperms and gymnosperms or evergreen and deciduous tree species. Functional traits explained some of the variation in drought responses between species (i.e. increased from 30 to 37% when wood density and specific leaf area were included). Tree species with denser wood and lower specific leaf area showed lower mortality responses. Our results illustrate the value of functional traits for understanding patterns of drought-induced tree mortality and suggest that mortality could become increasingly widespread in the future.

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In the northern Colorado Front Range, fire suppression during the 20th century is believed to have created a high hazard of catastrophic fire in ponderosa pine (Pinus ponderosa) forests. Since the early 1990s, resource managers have increased the use of prescribed fires to re-create fire regimes and forest structures similar to those of the pre-Euro-American settlement period in order both to reduce fire hazard and to improve forest health. To improve understanding of historical fire regimes, we conducted a study of fire history along an elevational gradient from ϳ1830 to 2800 m in ponderosa pine forests in the northern Front Range. Fire-scar dates were determined from 525 partial cross sections from living and dead trees at 41 sample sites. Fire frequencies and fire intervals were analyzed in relation to changes in human activities and interannual climatic variability as recorded in instrumental climatic records and tree-ring proxy records.Prior to modern fire suppression, the low elevation, open ponderosa pine forests of the northern Front Range were characterized by frequent surface fires, similar in frequency to many other ponderosa pine ecosystems in the West. In contrast, in higher elevation forests (above ϳ2400 m) where ponderosa pine is mixed with Douglas-fir (Pseudotsuga menziesii) and lodgepole pine (Pinus contorta), the fire regime was characterized by a much lower fire frequency and included extensive stand-replacing fires as well as surface fires. In the mid-1800s there was a marked increase in fire occurrence that can be related both to Euro-American settlement and increased climatic variability. This episode of increased fire left a legacy of dense, even-aged stands in higher elevation ponderosa pine forests, whereas increased stand densities in low elevation forests are attributed mainly to fire exclusion during the 20th century.Warmer and drier spring-summers, indicated in instrumental climatic records and in tree-ring proxy records of climate , are strongly associated with years of widespread fire. Years of widespread fire also tend to be preceded two to four years by wetter than average springs that increase the production of fine fuels. Alternation of wet and dry periods over time periods of 2-5 years is conducive to fire spread and is strongly linked to El Niñ o-Southern Oscillation (ENSO) events. The warm (El Niñ o) phase of ENSO is associated with greater moisture availability during spring that results in a peak of fire occurrence several years following El Niñ o events. Conversely, dry springs associated with La Niñ a events were followed by more widespread fire during the same year.The 1600-1920 fire-scar record indicates that individual years during which high percentages of the 41 sample sites synchronously recorded fire have occurred at least several times per century. The association of these years of widespread fire with very strong ENSO events demonstrates the importance of ENSO-related climatic variabililty in creating extreme fire hazard at a landscape scale.

JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact support@jstor.org.. Ecological Society of America is collaborating with JSTOR to digitize, preserve and extend access to Ecology.Abstract. Facilitation of tree establishment by nurse shrubs, which ameliorate otherwise unfavorable microenvironmental conditions, is a widely studied phenomenon. However, relatively little is known about how facilitative influences change in relation to interannual climatic variability. In northern Patagonia, Argentina, we examined influences of potential nurse shrubs on the establishment of the conifer Austrocedrus chilensis and assessed the significance of those influences to establishment during years of contrasting climate. We also experimentally investigated the effects of nurse shrubs and different water availability on tree seedling emergence and survival.A strong spatial association of Austrocedrus juveniles with shrubs, both beneath shrub canopies and near shrub canopies, indicates that shrubs favorably influence tree regeneration and that in some habitats and time periods nurse plants appear to be required for successful tree seedling establishment. Protection from direct sunlight was the main factor contributed by shrubs that enhanced the germination and survival of Austrocedrus. During the 1995-1996 experiment, no seedlings survived in the unwatered interspaces between shrubs, whereas maximal survival was obtained by watering seedlings at shaded sites.The results of this study indicate that in the Patagonian ecotone the strength of facilitative associations between shrubs and Austrocedrus juveniles closely tracks annual climatic variability. During extremely warm dry years, recruitment of Austrocedrus is nil with or without protection by nurse shrubs. During cool wet years, establishment may occur both beneath shrubs and in open interspaces; however, during average years, which are still years with substantial drought stress, establishment of Austrocedrus appears to require nurse shrubs.

Widespread synchronous wildfires driven by climatic variation, such as those that swept western North America during 1996, 2000, and 2002, can result in major environmental and societal impacts. Understanding relationships between continental-scale patterns of drought and modes of sea surface temperatures (SSTs) such as El Niñ o-Southern Oscillation (ENSO), Pacific Decadal Oscillation (PDO), and Atlantic Multidecadal Oscillation (AMO) may explain how interannual to multidecadal variability in SSTs drives fire at continental scales. We used local wildfire chronologies reconstructed from fire scars on tree rings across western North America and independent reconstructions of SST developed from tree-ring widths at other sites to examine the relationships of multicentury patterns of climate and fire synchrony. From 33,039 annually resolved fire-scar dates at 238 sites (the largest paleofire record yet assembled), we examined forest fires at regional and subcontinental scales. Since 1550 CE, drought and forest fires covaried across the West, but in a manner contingent on SST modes. During certain phases of ENSO and PDO, fire was synchronous within broad subregions and sometimes asynchronous among those regions. In contrast, fires were most commonly synchronous across the West during warm phases of the AMO. ENSO and PDO were the main drivers of high-frequency variation in fire (interannual to decadal), whereas the AMO conditionally changed the strength and spatial influence of ENSO and PDO on wildfire occurrence at multidecadal scales. A current warming trend in AMO suggests that we may expect an increase in widespread, synchronous fires across the western U.S. in coming decades.Atlantic Multidecadal Oscillation ͉ El Niñ o Southern Oscillation ͉ fire history network ͉ ocean warming ͉ Pacific Decadal Oscillation