Mediterranean forest ecosystems will be increasingly affected by hotter drought and more frequent and severe wildfire events in the future. However, little is known about the longer-term responses of these forests to multiple disturbances and the forests\' capacity to maintain ecosystem function. This is particularly so for below-ground organisms, which have received less attention than those above-ground, despite their essential contributions to forest function. We investigated rhizosphere microbial communities in a resprouting Eucalyptus marginata forest, southwestern Australia, that had experienced a severe wildfire four years previously, and a hotter drought eight years previously. Our aim was to understand how microbial communities are affected over longer-term trajectories by hotter drought and wildfire, singularly, and in combination. Fungal and bacterial DNA was extracted from soil samples, amplified, and subjected to high throughput sequencing. Richness, diversity, composition, and putative functional groups were then examined. We found a monotonic decrease in fungal, but not bacterial, richness and diversity with increasing disturbance with the greatest changes resulting from the combination of drought and wildfire. Overall fungal and bacterial community composition reflected a stronger effect of fire than drought, but the combination of both produced the greatest number of indicator taxa for fungi, and a significant negative effect on the abundance of several fungal functional groups. Key mycorrhizal fungi, fungal saprotrophs and fungal pathogens were found at lower proportions in sites affected by drought plus wildfire. Wildfire had a positive effect on bacterial hydrogen and bacterial nitrogen recyclers. Fungal community composition was positively correlated with live tree height. These results suggest that microbial communities, in particular key fungal functional groups, are highly responsive to wildfire following drought. Thus, a legacy of past climate conditions such as hotter drought can be important for mediating the responses of soil microbial communities to subsequent disturbance like wildfire.

Ongoing climatic change is threatening the survival of drought-sensitive tree species, such as silver fir (Abies alba). Drought-induced dieback had been previously explored in this conifer, although the role played by tree-level genetic diversity and its relationship with growth patterns and soil microsite conditions remained elusive. We used double digest restriction-site-associated DNA sequencing (ddRADseq) to describe different genetic characteristics of five silver fir forests in the Spanish Pyrenees, including declining and non-declining trees. Single nucleotide polymorphisms (SNPs) were used to investigate the relationships between genetics, dieback, intraspecific trait variation (functional dendrophenotypic traits and leaf traits), local bioclimatic conditions, and rhizosphere soil properties. While there were no noticeable genetic differences between declining and non-declining trees, genome-environment associations with selection signatures were abundant, suggesting a strong influence of climate, soil physicochemical properties, and soil microbial diversity on local adaptation. These results provide novel insights into how genetics and diverse environmental factors are interrelated and highlight the need to incorporate genetic data into silver fir forest dieback studies to gain a better understanding of local adaptation.

In recent decades, die-off events in Pinus sylvestris populations have increased. The causes of these phenomena, which are usually related to local and regional extreme hot droughts, have been extensively investigated from a physiological viewpoint. However, the consequences of die-off process in terms of demography and vegetation dynamics have been less thoroughly addressed. Here, we projected P. sylvestris plot dynamics after a die-off event, under climate change scenarios, considering also their early demographic stages (i.e., seedlings, saplings and ingrowth from the sapling to adult class), to assess the resilience of P. sylvestris populations after such events. We used Integral Projection Models (IPMs) to project future plot structure under current climate, and under RCP4.5 and RCP8.0 climate scenarios, using climatic suitability - extracted from Species Distribution Models - as a covariable in the estimations of vital rates over time. Field data feeding IPMs were obtained from two successive surveys, at the end of the die-off event (2013) and four years later (2017), undertaken on populations situated across the P. sylvestris range of distribution in Catalonia (NE Spain). Plots affected by die-off experienced a loss of large trees, which causes that basal area, tree diameter and tree density will remain lower for decades relative to unaffected plots. After the event, this situation is partially counterbalanced in affected plots by a greater increase in basal area and seedling recruitment into tree stage, thus promoting resilience. However, resilience is delayed under the climate-change scenarios with warmer and drier conditions involving additional physiological stress, due to a reduced abundance of seedlings and a smaller plot basal area. The study shows lagged effect of drought-induced die-off events on forest structure, also revealing stabilizing mechanisms, such as recruitment and tree growth release, which enhance resilience. However, these mechanisms would be jeopardized by oncoming regional warming.

Rear-edge populations at the xeric distribution limit of tree species are particularly vulnerable to forest dieback triggered by drought. This is the case of silver fir (Abies alba) forests located in Southwestern Europe. While silver fir drought-induced dieback patterns have been previously explored, information on the role played by nutritional impairment is lacking despite its potential interactions with tree carbon-water balances. We performed a comparative analysis of radial growth, intrinsic water-use efficiency (iWUE), oxygen isotopes (δ18 O) and nutrient concentrations in leaves of declining (DD) and non-declining (ND) trees in silver fir in four forests in the Spanish Pyrenees. We also evaluated the relationships among dieback predisposition, intraspecific trait variation (wood density and leaf traits) and rhizosphere soil physical-chemical properties. The onset of growth decline in DD trees occurred more than two decades ago, and they subsequently showed low growth resilience against droughts. The DD trees presented consistently lower foliar concentrations of nutrients such as P, K, Cu and Ni than ND trees. The strong effects of foliar nutrient status on growth resilience indices support the key role played by mineral nutrition in tree functioning and growth before, during and after drought. In contrast, variability in wood density and leaf morphological traits, as well as soil properties, showed weak relationships with tree nutritional status and drought performance. At the low elevation, warmer sites, DD trees showed stronger climate-growth relationships and lower δ18 O than ND trees. The uncoupling between iWUE and δ18 O, together with the positive correlations between P and K leaf concentrations and δ18 O, point to deeper soil/bedrock water sources and vertical decoupling between nutrient and water uptake in DD trees. This study provides novel insights into the mechanisms driving silver fir dieback and highlights the need to incorporate tree nutrition into forest dieback studies.
Las poblaciones del límite xérico de distribución de las especies de árboles son particularmente vulnerables al decaimiento forestal inducido por sequía. Este es el caso de los bosques de abeto (Abies alba) situados en el suroeste de Europa. Si bien los patrones de decaimiento provocado por sequía del abeto se han explorado previamente, falta información sobre el papel que desempeña el deterioro nutricional a pesar de sus interacciones potenciales con los balances de agua y carbono de los árboles. En este estudio, hemos realizado un análisis comparativo del crecimiento radial, la eficiencia intrínseca del uso del agua (iWUE), los isótopos de oxígeno (δ18O) y las concentraciones de nutrientes en hojas de árboles decaídos (DD) y no decaídos (ND) en cuatro abetares de los Pirineos españoles. También evaluamos las relaciones entre la predisposición al decaimiento, la variación de rasgos intraespecíficos (densidad de la madera y rasgos de las hojas) y las propiedades físico-químicas de la rizosfera. El inicio de la disminución del crecimiento en los árboles DD ocurrió hace más de dos décadas y posteriormente mostraron una baja resiliencia de crecimiento frente a las sequías. Los árboles DD presentaron concentraciones foliares consistentemente más bajas de nutrientes como P, K, Cu y Ni que los árboles ND. Los fuertes efectos del estado de los nutrientes foliares en los índices de resiliencia del crecimiento respaldan el papel clave que desempeña la nutrición mineral en el funcionamiento y el crecimiento de los árboles antes, durante y después de la sequía. En contraste, la variabilidad en la densidad de la madera y los rasgos morfológicos de las hojas, así como las propiedades del suelo, mostraron una relación débil con el estado nutricional de los árboles y la respuesta del crecimiento a la sequía. En los sitios más cálidos y de baja elevación, los árboles DD mostraron relaciones clima-crecimiento más fuertes y un δ18 O más bajo que los árboles ND. El desacoplamiento entre iWUE y δ18 O, junto con las correlaciones positivas entre las concentraciones foliares de P y K y δ18 O, apuntan a fuentes de agua más profundas del suelo/lecho rocoso y un desacoplamiento vertical entre la absorción de nutrientes y agua en los árboles DD. Este estudio proporciona información novedosa sobre los mecanismos que impulsan el decaimiento del abeto y destaca la necesidad de incorporar la nutrición de los árboles en los estudios de muerte regresiva del bosque.

Forest dieback processes linked to drought are expected to increase due to climate warming. Remotely sensed data offer several advantages over common field monitoring methods such as the ability to observe large areas on a systematic basis and monitoring their changes, making them increasingly used to assess changes in forest health. Here we aim to use a combined approximation of fieldwork and remote sensing to explore possible links between forest dieback and land surface phenological and trend variables derived from long Landsat time series. Forest dieback was evaluated in the field over 31 plots in a Mediterranean, xeric Pinus pinaster forest. Landsat 31-year time series of three greenness (EVI, NDVI, SAVI) and two wetness spectral indices (NMDI and TCW) were derived covering the period 1990-2020. Spectral indices from time series were decomposed into trend and seasonality using a Bayesian estimator while the relationships of the phenological and trend variables among levels of damage were assessed using linear and additive mixed models. We have not found any statistical pieces of evidence of extension or shortening patterns for the length of the phenological season over the examined 31-year period. Our results indicate that the dieback process was mainly related to the trend component of the spectral indices series whereas the phenological metrics were not related to forest dieback. We also found that plots with more dying or damaged trees displayed lower spectral indices trends after a severe drought event in the middle of the 1990s, which confirms the Landsat-derived spectral indices as indicators of early-warning signals. Drops in trends occurred earlier for wetness indices rather than for greenness indices which suggests that the former could be more appropriate for dieback detection, i.e. they could be used as early warning signals of impending loss of tree vigor.

In recent decades, many forest die-off events have been reported in relation to climate-change-induced episodes, such as droughts and heat waves. To understand how these extreme climatic events induce forest die-off, it is important to find a tool to standardize the climatic conditions experienced by different populations during a specific climatic event, taking into account the historic climatic conditions of the site where these populations live (bioclimatic niche). In this study, we used estimates of climatic suitability calculated from species distribution models (SDMs) for such purpose. We studied forest die-off across France during the 2003 heatwave that affected Western Europe, using 2,943 forest inventory plots dominated by 14 single tree species. Die-off severity was estimated by Normalized Difference Vegetation Index (NDVI) loss using Moderate-resolution Imaging Spectroradiometer remote sensor imagery. Climatic suitability at the local level during the historical 1979-2002 period (HCS), the episode time (2003; ECS) and suitability deviance during the historical period (HCS-SD) were calculated for each species by means of boosted regression tree models using the CHELSA climate database and occurrences extracted from European forest inventories. Low HCS-SD and high mean annual temperature explained the overall regional pattern of vulnerability to die-off across different monospecific forests. The combination of high historical and low episode climatic suitability also contributed significantly to overall forest die-off. Furthermore, we observed different species-specific relationships between die-off vulnerability and climatic suitability: Sub-Mediterranean and Mediterranean species tended to be vulnerable in historically more suitable localities (high HCS), whereas Euro-Siberian species presented greater vulnerability when the hot drought episode was more intense. We demonstrated that at regional scale, past climatic legacy plays an important role in explaining NDVI loss during the episode. Moreover, we demonstrated that SDMs-derived indexes, such as HCS, ECS and HCS-SD, could constitute a tool for standardizing the ways that populations and species experience climatic variability across time and space.

Extreme climatic and weather events are increasing in frequency and intensity across the world causing episodes of widespread tree mortality in many forested ecosystems. However, we have a limited understanding about which local factors influence tree mortality patterns, restricting our ability to predict tree mortality, especially within topographically complex tropical landscapes with a matrix of mature and secondary forests. We investigated the effects of two major local factors, topography and forest successional type, on climate-induced tropical tree mortality patterns using an observational and modeling approach. The northernmost Neotropical dry forest endured an unprecedented episode of frost-induced tree mortality after the historic February 2011 cold wave hit northwestern Mexico. In a moderately hilly landscape covering mature and secondary tropical dry forests, we surveyed 454 sites for the presence or absence of frost-induced tree mortality. In addition, across forty-eight 1 ha plots equally split into the two forest types, we examined 6,981 woody plants to estimate a frost-disturbance severity metric using the density of frost-killed trees. Elevation is the main factor modulating frost effects regardless of forest type. Higher occurrence probabilities of frost-induced tree mortality at lowland forests can be explained by the strong influence of elevation on temperature distribution since heavier cold air masses move downhill during advective frosts. Holding elevation constant, the probability of frost-induced tree mortality in mature forests was twice that of secondary forests but severity showed the opposite pattern, suggesting a cautious use of occurrence probabilities of tree mortality to infer severity of climate-driven disturbances. Extreme frost events, in addition to altering forest successional pathways and ecosystem services, likely maintain and could ultimately shift latitudinal and altitudinal range margins of Neotropical dry forests.

Globally, trees are increasingly dying from extreme drought, a trend that is expected to increase with climate change. Loss of trees has significant ecological, biophysical, and biogeochemical consequences. In 2011, a record drought caused widespread tree mortality in Texas. Using remotely sensed imagery, we quantified canopy loss during and after the drought across the state at 30-m spatial resolution, from the eastern pine/hardwood forests to the western shrublands, a region that includes the boundaries of many species ranges. Canopy loss observations in ~200 multitemporal fine-scale orthophotos (1-m) were used to train coarser Landsat imagery (30-m) to create 30-m binary statewide canopy loss maps. We found that canopy loss occurred across all major ecoregions of Texas, with an average loss of 9.5%. The drought had the highest impact in post oak woodlands, pinyon-juniper shrublands and Ashe juniper woodlands. Focusing on a 100-km by ~1,000-km transect spanning the State\'s fivefold east-west precipitation gradient (~1,500 to ~300 mm), we compared spatially explicit 2011 climatic anomalies to our canopy loss maps. Much of the canopy loss occurred in areas that passed specific climatic thresholds: warm season anomalies in mean temperature (+1.6°C) and vapor pressure deficit (VPD, +0.66 kPa), annual percent deviation in precipitation (-38%), and 2011 difference between precipitation and potential evapotranspiration (-1,206 mm). Although similarly low precipitation occurred during the landmark 1950s drought, the VPD and temperature anomalies observed in 2011 were even greater. Furthermore, future climate data under the representative concentration pathway 8.5 trajectory project that average values will surpass the 2011 VPD anomaly during the 2070-2099 period and the temperature anomaly during the 2040-2099 period. Identifying vulnerable ecological systems to drought stress and climate thresholds associated with canopy loss will aid in predicting how forests will respond to a changing climate and how ecological landscapes will change in the near term.

Increases in drought and temperature stress in forest and woodland ecosystems are thought to be responsible for the rise in episodic mortality events observed globally. However, key climatic drivers common to mortality events and the impacts of future extreme droughts on tree survival have not been evaluated. Here, we characterize climatic drivers associated with documented tree die-off events across Australia using standardized climatic indices to represent the key dimensions of drought stress for a range of vegetation types. We identify a common probabilistic threshold associated with an increased risk of die-off across all the sites that we examined. We show that observed die-off events occur when water deficits and maximum temperatures are high and exist outside 98% of the observed range in drought intensity; this threshold was evident at all sites regardless of vegetation type and climate. The observed die-off events also coincided with at least one heat wave (three consecutive days above the 90th percentile for maximum temperature), emphasizing a pivotal role of heat stress in amplifying tree die-off and mortality processes. The joint drought intensity and maximum temperature distributions were modeled for each site to describe the co-occurrence of both hot and dry conditions and evaluate future shifts in climatic thresholds associated with the die-off events. Under a relatively dry and moderate warming scenario, the frequency of droughts capable of inducing significant tree die-off across Australia could increase from 1 in 24 years to 1 in 15 years by 2050, accompanied by a doubling in the occurrence of associated heat waves. By defining commonalities in drought conditions capable of inducing tree die-off, we show a strong interactive effect of water and high temperature stress and provide a consistent approach for assessing changes in the exposure of ecosystems to extreme drought events.

Forests around the world are subject to risk of high rates of tree growth decline and increased tree mortality from combinations of climate warming and drought, notably in semi-arid settings. Here, we assess how climate warming has affected tree growth in one of the world\'s most extensive zones of semi-arid forests, in Inner Asia, a region where lack of data limits our understanding of how climate change may impact forests. We show that pervasive tree growth declines since 1994 in Inner Asia have been confined to semi-arid forests, where growing season water stress has been rising due to warming-induced increases in atmospheric moisture demand. A causal link between increasing drought and declining growth at semi-arid sites is corroborated by correlation analyses comparing annual climate data to records of tree-ring widths. These ring-width records tend to be substantially more sensitive to drought variability at semi-arid sites than at semi-humid sites. Fire occurrence and insect/pathogen attacks have increased in tandem with the most recent (2007-2009) documented episode of tree mortality. If warming in Inner Asia continues, further increases in forest stress and tree mortality could be expected, potentially driving the eventual regional loss of current semi-arid forests.