Under Stress: Searching for Genes Involved in the Response of Abies pinsapo Boiss to Climate Change.

Currently, Mediterranean forests are experiencing the deleterious effects of global warming, which mainly include increased temperatures and decreased precipitation in the region. Relict Abies pinsapo fir forests, endemic in the southern Iberian Peninsula, are especially sensitive to these recent environmental disturbances, and identifying the genes involved in the response of this endangered tree species to climate-driven stresses is of paramount importance for mitigating their effects. Genomic resources for A. pinsapo allow for the analysis of candidate genes reacting to warming and aridity in their natural habitats. Several members of the complex gene families encoding late embryogenesis abundant proteins (LEAs) and heat shock proteins (HSPs) have been found to exhibit differential expression patterns between wet and dry seasons when samples from distinct geographical locations and dissimilar exposures to the effects of climate change were analyzed. The observed changes were more perceptible in the roots of trees, particularly in declining forests distributed at lower altitudes in the more vulnerable mountains. These findings align with previous studies and lay the groundwork for further research on the molecular level. Molecular and genomic approaches offer valuable insights for mitigating climate stress and safeguarding this endangered conifer.

Sapwood mycobiome varies across host, plant compartment and environments in Nothofagus forests from Northern Patagonia.

Global forests are increasingly being threatened by altered climatic conditions and increased attacks by pests and pathogens. The complex ecological interactions among pathogens, microbial communities, tree hosts and the environment are important drivers of forest dynamics. Little is known about the ecology of forest pathology and related microbial communities in temperate forests of the southern hemisphere. In this study, we used next-generation sequencing to characterize sapwood-inhabiting fungal communities in North Patagonian Nothofagus forests and assessed patterns of diversity of taxa and ecological guilds across climatic, site and host variables (health condition and compartment) as a contribution to Nothofagus autecology. The diversity patterns inferred through the metabarcoding analysis were similar to those obtained through culture-dependent approaches. However, we detected additional heterogeneity and greater richness with culture-free methods. Host species was the strongest driver of fungal community structure and composition, while host health status was the weakest. The relative impacts of site, season, plant compartment and health status were different for each tree species; these differences can be interpreted as a matter of water availability. For Nothofagus dombeyi, which is distributed across a wide range of climatic conditions, site was the strongest driver of community composition. The microbiome of N. pumilio varied more with season and temperature, a relevant factor for forest conservation in the present climate change scenario. Both species carry a number of potential fungal pathogens in their sapwood, whether they exhibit symptoms or not. Our results provide insight into the diversity of fungi associated with the complex pathobiome of the dominant Nothofagus species in southern South America.

Los bosques del mundo están cada vez más amenazados por las condiciones climáticas alteradas y el aumento de los ataques de plagas y patógenos. Las complejas interacciones ecológicas entre los patógenos, las comunidades microbianas, los árboles hospedantes y el medio ambiente son impulsores importantes de la dinámica forestal. Poco se sabe sobre la ecología de la patología forestal y las comunidades microbianas relacionadas en los bosques templados del hemisferio sur. En este estudio, utilizamos la secuenciación Illumina para caracterizar las comunidades de hongos que habitan en la albura en los bosques de Nothofagus de la Patagonia Norte y evaluamos los patrones de diversidad de taxones y gremios ecológicos a través de variables climáticas, de sitio y de hospedante (identidad, condición de salud y compartimento) como una contribución a la autoecología de los Nothofagus. Los patrones de diversidad inferidos a través del análisis metabarcoding fueron similares a los obtenidos a través de enfoques dependientes de cultivo. Sin embargo, detectamos mayor heterogeneidad y mayor riqueza con métodos independientes de cultivo. La especie hospedante fue el modelador más fuerte de la estructura y composición de la comunidad fúngica, mientras que el estado de salud del hospedante fue el más débil. El impacto relativo del sitio, la estación, el compartimento y el estado de salud fueron diferentes para cada especie de árbol; estas diferencias pueden interpretarse en clave de disponibilidad de agua. Para N. dombeyi, que se distribuye a lo largo de una amplia gama de condiciones climáticas, el sitio fue el principal modelador de la composición de la comunidad. El micobioma de Nothofagus pumilio varió más con la estación y la temperatura, un factor relevante para la conservación de los bosques en el escenario actual de cambio climático. Ambas especies portan una serie de patógenos fúngicos potenciales en su albura, ya sea que muestren síntomas o no. Nuestros resultados brindan una idea de la diversidad de hongos asociados con el complejo patobioma de las especies dominantes de Nothofagus en el sur de América del Sur.

Climate-driven tree growth and mortality in the Black Forest, Germany-Long-term observations.

Episodic tree mortality can be caused by various reasons. This study describes climate-driven tree mortality and tree growth in the Black Forest mountain range in Germany. It is based on a 68-year consistent data series describing the annual mortality of all trees growing in a forest area of almost 250 thousand ha. The study excludes mortality caused by storm, snow and ice, and fire. The sequence of the remaining mortality, the so-called "desiccated trees," is analyzed and compared with the sequence of the climatic water balance during the growing season and the annual radial growth of Norway spruce in the Black Forest. The annual radial growth series covers 121 years and the climatic water balance series 140 years. These unique time series enable a quantitative assessment of multidecadal drought and heat impacts on growth and mortality of forest trees on a regional spatial scale. Data compiled here suggest that the mortality of desiccated trees in the Black Forest during the last 68 years is driven by the climatic water balance. Decreasing climatic water balance coincided with an increase in tree mortality and growth decline. Consecutive hot and dry summers enhance mortality and growth decline as a consequence of drought legacies lasting several years. The sensitivity of tree growth and mortality to changes in the climatic water balance increases with the decreasing trend of the climatic water balance. The findings identify the climatic water balance as the main driver of mortality and growth variation during the 68-year observation period on a landscape-scale including a variety of different sites. They suggest that bark beetle population dynamics modify mortality rates. They as well provide evidence that the mortality during the last 140 years never was as high as in the most recent years.

First Report of Dieback of Quercus suber Trees Associated with Phytophthora quercina in Morocco.

Cork oak (Quercus suber L.) is an evergreen tree native to SW Europe and NW Africa. It covers 2·106 ha in the western Mediterranean basin, forms heterogeneous forest ecosystems and represents an important source of income derived from cork production. While in Iberia, Italy, Tunisia and Algeria, drought and several endemic pathogens have been associated with cork oak decline (Moricca et al. 2016; Smahi et al. 2017), in Morocco there is no evidence, apart from overgrazing and human intervention (Fennane and Rejdali 2015), of a pathogen associated with oak decline. In December 2019, extensive dieback and mortality of 60-year-old cork oak trees were observed in a natural stand of ca 150 ha located 5 km east from Touazithe, in Maâmora forest, Morocco (34°13'38''N, 6°14'51''W - 87 m a.s.l.). Two years before, Q. suber seedlings from a local nursery were planted to increase tree density. Symptoms in trees and planted seedlings included chlorosis, reddish-brown discoloration of the whole crown and dieback starting in the upper crown. Root rot and lack of fine roots were observed. Tree mortality was estimated at ca 30%, and disease incidences of trees and seedlings were 45 and 70%, respectively. A Phytophthora species was consistently isolated from the rhizosphere of 3 symptomatic trees randomly selected at the site using leaves as bait (Jung et al. 1996). On carrot agar Phytophthora colonies were uniform and cottonwool-like. Sporangia were typically terminal, with ovoid, and obpyriform shape, mostly papillate, measuring 30.7 ± 4.7 µm length and 22.7 ± 4.1 µm wide. Oogonia were produced in single culture, and they were globose to subglobose, elongated to ellipsoid, 32.1 ± 2.9 µm in diameter and 46.1 ± 4.8 µm in length. Oospores were usually spherical, thick-walled, and measured 28.1 ± 2.4 µm. Antheridia were paragynous, mostly spherical, measuring 12.2 ± 1.4 µm. Isolates had minimum and maximum temperatures of 5 °C and 30 °C, respectively, and a growth optimum at 20 °C. Apart from the small size of sporangia, features were typical of Phytophthora quercina Jung. The identity of a representative strain (TJ1500) was corroborated by sequencing the ITS and mitochondrial cox1 gene regions, and BLAST search in GenBank showed 100% homology with sequences of the ex-type culture of P. quercina (KF358229 and KF358241 accessions, respectively). Both sequences of the representative isolate were submitted to GenBank (accessions OP086243 and OP290549). The strain TJ1500 is currently stored within the culture collections of the Mendel University in Brno and the University of Sassari. Its pathogenicity was verified and compared with a P. cinnamomi strain in a soil infestation test with one-year-old cork oak seedlings (Corcobado et al. 2017). Five months after inoculation, the symptoms described were observed in the seedlings, and fine root weight of plants inoculated with the TJ1500 strain and P. cinnamomi was reduced by 19 and 42%, respectively, in relation to non-inoculated controls. The pathogen was re-isolated from the necrotic roots, thus fulfilling Koch's postulates. So far, P. quercina has been reported associated with chronic mortality of cork oak in new plantations in Spain (Martín-García et al. 2015; Jung et al. 2016) and natural forests in Italy (Seddaiu et al. 2020). To our knowledge this is the first report of P. quercina in Morocco. Givenat Morocco is an important cork producing country, our finding warns about the risk this pathogen poses to Q. suber and other North African oaks.

Chlorophyll content estimation in an open-canopy conifer forest with Sentinel-2A and hyperspectral imagery in the context of forest decline.

With the advent of Sentinel-2, it is now possible to generate large-scale chlorophyll content maps with unprecedented spatial and temporal resolution, suitable for monitoring ecological processes such as vegetative stress and/or decline. However methodological gaps exist for adapting this technology to heterogeneous natural vegetation and for transferring it among vegetation species or plan functional types. In this study, we investigated the use of Sentinel-2A imagery for estimating needle chlorophyll (Ca+b) in a sparse pine forest undergoing significant needle loss and tree mortality. Sentinel-2A scenes were acquired under two extreme viewing geometries (June vs. December 2016) coincident with the acquisition of high-spatial resolution hyperspectral imagery, and field measurements of needle chlorophyll content and crown leaf area index. Using the high-resolution hyperspectral scenes acquired over 61 validation sites we found the CI chlorophyll index R750/R710 and Macc index (which uses spectral bands centered at 680 nm, 710 nm and 780 nm) had the strongest relationship with needle chlorophyll content from individual tree crowns (r2 = 0.61 and r2 = 0.59, respectively; p < 0.001), while TCARI and TCARI/OSAVI, originally designed for uniform agricultural canopies, did not perform as well (r2 = 0.21 and r2 = 0.01, respectively). Using lower-resolution Sentinel-2A data validated against hyperspectral estimates and ground truth needle chlorophyll content, the red-edge index CI and the Sentinel-specific chlorophyll indices CI-Gitelson, NDRE1 and NDRE2 had the highest accuracy (with r2 values >0.7 for June and >0.4 for December; p < 0.001). The retrieval of needle chlorophyll content from the entire Sentinel-2A bandset using the radiative transfer model INFORM yielded r2 = 0.71 (RMSE = 8.1 µg/cm2) for June, r2 = 0.42 (RMSE = 12.2 µg/cm2) for December, and r2 = 0.6 (RMSE = 10.5 µg/cm2) as overall performance using the June and December datasets together. This study demonstrates the retrieval of leaf Ca+b with Sentinel-2A imagery by red-edge indices and by an inversion method based on a hybrid canopy reflectance model that accounts for tree density, background and shadow components common in sparse forest canopies.

Hydraulics play an important role in causing low growth rate and dieback of aging Pinus sylvestris var. mongolica trees in plantations of Northeast China.

The frequently observed forest decline in water-limited regions may be associated with impaired tree hydraulics, but the precise physiological mechanisms remain poorly understood. We compared hydraulic architecture of Mongolian pine (Pinus sylvestris var. mongolica) trees of different size classes from a plantation and a natural forest site to test whether greater hydraulic limitation with increasing size plays an important role in tree decline observed in the more water-limited plantation site. We found that trees from plantations overall showed significantly lower stem hydraulic efficiency. More importantly, plantation-grown trees showed significant declines in stem hydraulic conductivity and hydraulic safety margins as well as syndromes of stronger drought stress with increasing size, whereas no such trends were observed at the natural forest site. Most notably, the leaf to sapwood area ratio (LA/SA) showed a strong linear decline with increasing tree size at the plantation site. Although compensatory adjustments in LA/SA may mitigate the effect of increased water stress in larger trees, they may result in greater risk of carbon imbalance, eventually limiting tree growth at the plantation site. Our results provide a potential mechanistic explanation for the widespread decline of Mongolian pine trees in plantations of Northern China.

Contributions of insects and droughts to growth decline of trembling aspen mixed boreal forest of western Canada.

Insects, diseases, fire and drought and other disturbances associated with global climate change contribute to forest decline and mortality in many parts of the world. Forest decline and mortality related to drought or insect outbreaks have been observed in North American aspen forests. However, little research has been done to partition and estimate their relative contributions to growth declines. In this study, we combined tree-ring width and basal area increment series from 40 trembling aspen (Populus tremuloides Michx.) sites along a latitudinal gradient (from 52° to 58°N) in western Canada and attempted to investigate the effect of drought and insect outbreaks on growth decline, and simultaneously partition and quantify their relative contributions. Results indicated that the influence of drought on forest decline was stronger than insect outbreaks, although both had significant effects. Furthermore, the influence of drought and insect outbreaks showed spatiotemporal variability. In addition, our data suggest that insect outbreaks could be triggered by warmer early spring temperature instead of drought, implicating that potentially increased insect outbreaks are expected with continued warming springs, which may further exacerbate growth decline and death in North America aspen mixed forests.

Alterations in leaf nitrogen metabolism indicated the structural changes of subtropical forest by canopy addition of nitrogen.

Globally, nitrogen deposition increment has caused forest structural changes due to imbalanced plant nitrogen metabolism and subsequent carbon assimilation. Here, a 2 consecutive-year experiment was conducted to reveal the effects of canopy addition of nitrogen (CAN) on nitrogen absorption, assimilation, and allocation in leaves of three subtropical forest woody species (Castanea henryi, Ardisia quinquegona, and Blastus cochinchinensis). We hypothesized that CAN altered leaf nitrogen absorption, assimilation and partitioning of different plants in different ways in subtropical forest. It shows that CAN increased maximum photosynthetic rate (Amax), photosynthetic nitrogen use efficiency (PNUE), and metabolic protein content of the two understory species A. quinquegona and B. cochinchinensis. By contrary, for the overstory species, C. henryi, Amax, PNUE, and metabolic protein content were significantly reduced in response to CAN. We found that changes in leaf nitrogen metabolism were mainly due to the differences in enzyme (e.g. Ribulose-1,5-bisphosphate carboxylase, nitrate reductase, nitrite reductase and glutamine synthetase) activities under CAN treatment. Our results indicated that C. henryi may be more susceptible to CAN treatment, and both A. quinquegona and B. cochinchinensis could better adapt to CAN treatment but in different ways. Our findings may partially explain the ongoing degradation of subtropical forest into a community dominated by small trees and shrubs in recent decades. It is possible that persistent high levels of atmospheric nitrogen deposition will lead to the steady replacement of dominant woody species in this subtropical forest.

Understanding the temporal dimension of the red-edge spectral region for forest decline detection using high-resolution hyperspectral and Sentinel-2a imagery.

The operational monitoring of forest decline requires the development of remote sensing methods that are sensitive to the spatiotemporal variations of pigment degradation and canopy defoliation. In this context, the red-edge spectral region (RESR) was proposed in the past due to its combined sensitivity to chlorophyll content and leaf area variation. In this study, the temporal dimension of the RESR was evaluated as a function of forest decline using a radiative transfer method with the PROSPECT and 3D FLIGHT models. These models were used to generate synthetic pine stands simulating decline and recovery processes over time and explore the temporal rate of change of the red-edge chlorophyll index (CI) as compared to the trajectories obtained for the structure-related Normalized Difference Vegetation Index (NDVI). The temporal trend method proposed here consisted of using synthetic spectra to calculate the theoretical boundaries of the subspace for healthy and declining pine trees in the temporal domain, defined by CItime=n/CItime=n+1 vs. NDVItime=n/NDVItime=n+1. Within these boundaries, trees undergoing decline and recovery processes showed different trajectories through this subspace. The method was then validated using three high-resolution airborne hyperspectral images acquired at 40 cm resolution and 260 spectral bands of 6.5 nm full-width half-maximum (FWHM) over a forest with widespread tree decline, along with field-based monitoring of chlorosis and defoliation (i.e., 'decline' status) in 663 trees between the years 2015 and 2016. The temporal rate of change of chlorophyll vs. structural indices, based on reflectance spectra extracted from the hyperspectral images, was different for trees undergoing decline, and aligned towards the decline baseline established using the radiative transfer models. By contrast, healthy trees over time aligned towards the theoretically obtained healthy baseline. The applicability of this temporal trend method to the red-edge bands of the MultiSpectral Imager (MSI) instrument on board Sentinel-2a for operational forest status monitoring was also explored by comparing the temporal rate of change of the Sentinel-2-derived CI over areas with declining and healthy trees. Results demonstrated that the Sentinel-2a red-edge region was sensitive to the temporal dimension of forest condition, as the relationships obtained for pixels in healthy condition deviated from those of pixels undergoing decline.

Drought-induced shoot dieback starts with massive root xylem embolism and variable depletion of nonstructural carbohydrates in seedlings of two tree species.

Combining hydraulic- and carbon-related measurements helps to understand drought-induced plant mortality. Here, we investigated the role that plant respiration (R) plays in determining carbon budgets under drought. We measured the hydraulic conductivity of stems and roots, and gas exchange and nonstructural carbohydrate (NSC) concentrations of leaves, stems and roots of seedlings of two resprouting species exposed to drought or well-watered conditions: Ulmus minor (riparian tree) and Quercus ilex (dryland tree). With increasing water stress (occurring more rapidly in larger U. minor), declines in leaf, stem and root R were less pronounced than that in leaf net photosynthetic CO2 uptake (Pn ). Daytime whole-plant carbon gain was negative below -4 and -6 MPa midday xylem water potential in U. minor and Q. ilex, respectively. Relative to controls, seedlings exhibiting shoot dieback suffered c. 80% loss of hydraulic conductivity in both species, and reductions in NSC concentrations in U. minor. Higher drought-induced depletion of NSC reserves in U. minor was related to higher plant R, faster stomatal closure, and premature leaf-shedding. Differences in drought resistance relied on the ability to maintain hydraulic conductivity during drought, rather than tolerating conductivity loss. Root hydraulic failure elicited shoot dieback and precluded resprouting without root NSC reserves being apparently limiting for R.

Plant-soil feedbacks in declining forests: implications for species coexistence.

Plant-soil feedbacks (PSFs) play a relevant role as drivers of species abundance, coexistence, and succession in plant communities. However, the potential contribution of PSFs to community dynamics in changing forest ecosystems affected by global change drivers is still largely unexplored. We measured the direction, strength and nature (biological vs. chemical) of PSFs experienced by coexisting tree species in two types of declining Quercus suber forests of southwestern Spain (open woodland vs. closed forest) invaded by the exotic soil pathogen Phytophthora cinnamomi. To test PSFs in a realistic community context, we focused not only on individual PSFs (i.e., comparing the growth of a tree species on conspecific vs. heterospecific soil) but also calculated net-pairwise PSFs by comparing performance of coexisting tree species on their own and each other's soils. We hypothesized that the decline and death of Q. suber would alter the direction and strength of individual and net-pairwise PSFs due to the associated changes in soil nutrients and microbial communities, with implications for recruitment dynamics and species coexistence. In support of our hypothesis, we found that the decline of Q. suber translated into substantial alterations of individual and net-pairwise PSFs, which shifted from mostly neutral to significantly positive or negative, depending on the forest type. In both cases however the identified PSFs benefited other species more than Q. suber (i.e., heterospecific positive PSF in the open woodland, conspecific negative PSF in the closed forest). Our results supported PSFs driven by changes in chemical soil properties (mainly phosphorus) and arbuscular mycorrhizal fungi, but not in pathogen abundance. Overall, our study suggests that PSFs might reinforce the loss of dominance of Q. suber in declining forests invaded by P. cinnamomi by promoting the relative performance of non-declining coexisting species. More generally, our results indicate an increase in the strength of net PSFs as natural forests become disturbed by global change drivers (e.g., invasive species), suggesting an increasingly important role of PSFs in forest community dynamics in the near future.

Climate variability drives recent tree mortality in Europe.

Tree mortality is an important process in forest ecosystems, frequently hypothesized to be highly climate sensitive. Yet, tree death remains one of the least understood processes of forest dynamics. Recently, changes in tree mortality have been observed in forests around the globe, which could profoundly affect ecosystem functioning and services provisioning to society. We describe continental-scale patterns of recent tree mortality from the only consistent pan-European forest monitoring network, identifying recent mortality hotspots in southern and northern Europe. Analyzing 925,462 annual observations of 235,895 trees between 2000 and 2012, we determine the influence of climate variability and tree age on interannual variation in tree mortality using Cox proportional hazard models. Warm summers as well as high seasonal variability in precipitation increased the likelihood of tree death. However, our data also suggest that reduced cold-induced mortality could compensate increased mortality related to peak temperatures in a warming climate. Besides climate variability, age was an important driver of tree mortality, with individual mortality probability decreasing with age over the first century of a trees life. A considerable portion of the observed variation in tree mortality could be explained by satellite-derived net primary productivity, suggesting that widely available remote sensing products can be used as an early warning indicator of widespread tree mortality. Our findings advance the understanding of patterns of large-scale tree mortality by demonstrating the influence of seasonal and diurnal climate variation, and highlight the potential of state-of-the-art remote sensing to anticipate an increased likelihood of tree mortality in space and time.

Novel forest decline triggered by multiple interactions among climate, an introduced pathogen and bark beetles.

Novel forest decline is increasing due to global environmental change, yet the causal factors and their interactions remain poorly understood. Using tree ring analyses, we show how climate and multiple biotic factors caused the decline of whitebark pine (Pinus albicaulis) in 16 stands in the southern Canadian Rockies. In our study area, 72% of whitebark pines were dead and 18% had partially dead crowns. Tree mortality peaked in the 1970s; however, the annual basal area increment of disturbed trees began to decline significantly in the late 1940s. Growth decline persisted up to 30 years before trees died from mountain pine beetle (Dendroctonus ponderosae), Ips spp. bark beetles or non-native blister rust pathogen (Cronartium ribicola). Climate-growth relations varied over time and differed among the healthy and disturbed subpopulations of whitebark pine. Prior to the 1940s, cool temperatures limited the growth of all subpopulations. Growth of live, healthy trees became limited by drought during the cool phase (1947 -1976) of the Pacific Decadal Oscillation (PDO) and then reverted to positive correlations with temperature during the subsequent warm PDO phase. In the 1940s, the climate-growth relations of the disturbed subpopulations diverged from the live, healthy trees with trees ultimately killed by mountain pine beetle diverging the most. We propose that multiple factors interacted over several decades to cause unprecedented rates of whitebark pine mortality. Climatic variation during the cool PDO phase caused drought stress that may have predisposed trees to blister rust. Subsequent decline in snowpack and warming temperatures likely incited further climatic stress and with blister rust reduced tree resistance to bark beetles. Ultimately, bark beetles and blister rust contributed to tree death. Our findings suggest the complexity of whitebark pine decline and the importance of considering multiway drought-disease-insect interactions over various timescales when interpreting forest decline.

Viewing forests from below: fine root mass declines relative to leaf area in aging lodgepole pine stands.

In the continued quest to explain the decline in productivity and vigor with aging forest stands, the most poorly studied area relates to root system change in time. This paper measures the wood production, root and leaf area (and mass) in a chronosequence of fire-origin lodgepole pine (Pinus contorta Loudon) stands consisting of four age classes (12, 21, 53, and ≥100 years), each replicated ~ five times. Wood productivity was greatest in the 53-year-old stands and then declined in the ≥100-year-old stands. Growth efficiency, the quantity of wood produced per unit leaf mass, steadily declined with age. Leaf mass and fine root mass plateaued between the 53- and ≥100-year-old stands, but leaf area index actually increased in the older stands. An increase in the leaf area index:fine root area ratio supports the idea that older stand are potentially limited by soil resources. Other factors contributing to slower growth in older stands might be lower soil temperatures and increased self-shading due to the clumped nature of crowns. Collectively, the proportionally greater reduction in fine roots in older stands might be the variable that predisposes these forests to be at a potentially greater risk of stress-induced mortality.

Thinning effects on litterfall remaining after 8 years and improved stand resilience in Aleppo pine afforestation (SE Spain).

Monthly litterfall was monitored over a 3-year period in afforested Aleppo pines in the Mediterranean semiarid SE Spain with the aim of determining the long-term response of pines to reductions in tree competition and how this forest practice might influence stand resilience. Three thinning intensities applied 5 years earlier were evaluated (T75 = 75% of the basal area removed, T60 = 60% and T48 = 48%), both at the stand and at the tree level. On average, the total annual litterfall varied between 1.30 Mg ha(-1) yr(-1) (±0.24 SE) in T75 and 3.28 Mg ha(-1) yr(-1) (±0.78 SE) in the unthinned control. At the stand level, monthly differences among the treatments were found over time in the needles (F = 11.09, df = 3, P = 0.0009) and woody fraction (F = 4.36, df = 3, P = 0.0269) following the thinning gradient: T0 (control)>T48 > T60 > T75, and for the total amount of needles (χ(2) = 9.33, P = 0.025) and twigs (χ(2) = 9.11, P = 0.027) recorded at the end of the study period. High amounts of twig and needle fall were recorded during summer and beginning of autumn, whereas the main miscellanea inputs were registered during the spring, coinciding with the fall of nests and frass from caterpillar outbreaks. At the tree level, the total litterfall fluctuated between 1.5 kg tree yr(-1) in T0 (2nd yr) and 7.0 kg tree yr(-1) in T75 (3rd yr), although mean annual statistical differences among the treatments were found only for the first year of monitoring. However, needle fall was higher for larger pines (T75) than for the smaller ones in control (T0) when the data were analysed over the 3-year-period (F = 3.64, df = 3, P = 0.0247), and the same happened for the woody fraction (F = 3.63, df = 3, P = 0.0250). By contrast, pine trees in the unthinned control registered needle-fall rates (measured as kg m(-2) tree(-1)) that were similar to or higher than those of pine trees in thinned stands, suggesting that defoliation processes took place at high tree densities, especially after a severely dry period. We propose thinning as a measure to adapt high-density plantations to alterations due to climate change, in order to prevent forest decline and mortality.

A climate change context for the decline of a foundation tree species in south-western Australia: insights from phylogeography and species distribution modelling.

BACKGROUND AND AIMS: A worldwide increase in tree decline and mortality has been linked to climate change and, where these represent foundation species, this can have important implications for ecosystem functions. This study tests a combined approach of phylogeographic analysis and species distribution modelling to provide a climate change context for an observed decline in crown health and an increase in mortality in Eucalyptus wandoo, an endemic tree of south-western Australia. METHODS: Phylogeographic analyses were undertaken using restriction fragment length polymorphism analysis of chloroplast DNA in 26 populations across the species distribution. Parsimony analysis of haplotype relationships was conducted, a haplotype network was prepared, and haplotype and nucleotide diversity were calculated. Species distribution modelling was undertaken using Maxent models based on extant species occurrences and projected to climate models of the last glacial maximum (LGM). KEY RESULTS: A structured pattern of diversity was identified, with the presence of two groups that followed a climatic gradient from mesic to semi-arid regions. Most populations were represented by a single haplotype, but many haplotypes were shared among populations, with some having widespread distributions. A putative refugial area with high haplotype diversity was identified at the centre of the species distribution. Species distribution modelling showed high climatic suitability at the LGM and high climatic stability in the central region where higher genetic diversity was found, and low suitability elsewhere, consistent with a pattern of range contraction. CONCLUSIONS: Combination of phylogeography and paleo-distribution modelling can provide an evolutionary context for climate-driven tree decline, as both can be used to cross-validate evidence for refugia and contraction under harsh climatic conditions. This approach identified a central refugial area in the test species E. wandoo, with more recent expansion into peripheral areas from where it had contracted at the LGM. This signature of contraction from lower rainfall areas is consistent with current observations of decline on the semi-arid margin of the range, and indicates low capacity to tolerate forecast climatic change. Identification of a paleo-historical context for current tree decline enables conservation interventions to focus on maintaining genetic diversity, which provides the evolutionary potential for adaptation to climate change.

A 10-Year Assessment of Hemlock Decline in the Catskill Mountain Region of New York State Using Hyperspectral Remote Sensing Techniques.

The hemlock woolly adelgid is a serious pest of Eastern and Carolina hemlock in the eastern United States. Successfully managing the hemlock resource in the region depends on careful monitoring of the spread of this invasive pest and the targeted application of management options such as biological control, chemical, or silvicultural treatments. To inform these management activities and test the applicability of a landscape-scale remote sensing effort to monitor hemlock condition, hyperspectral collections, and concurrent ground-truthing in 2001 and 2012 of hemlock condition were compared with field metrics spanning a 10-yr survey in the Catskills region of New York. Fine twig dieback significantly increased from 9 to 15% and live crown ratio significantly decreased from 67 to 56% in 2001 and 2012, respectively. We found a significant shift from 59% "healthy" hemlock in 2001 to only 16% in 2012. However, this shift from healthy to declining classifications was mostly a shift to decline class 2 "early decline". These results indicate that while there has been significant increase in decline symptoms as measured in both field and remote sensing assessments, a majority of the declining areas identified in the resulting spatial coverages remain in the "early decline" category and widespread mortality has not yet occurred. While this slow decline across the region stands in contrast to many reports of mortality within 10 yr, the results from this work are in line with other long-term monitoring studies and indicate that armed with the spatial information provided here, continued management strategies can be focused on particular areas to help control the further decline of hemlock in the region.

First Report of Diplodia quercivora and Neofusicoccum vitifusiforme Associated with Cankers and Necrosis of Holm Oak (Quercus ilex) in Declining Stands in Southern Italy.

The emergence of new plant diseases is an increasingly important concern. Climate change is likely to be among the factors causing most of the emerging diseases endangering forest and tree heritage around the world. Such diseases may be caused by latent pathogens or microorganisms cryptically associated with plants. The shift from a non-pathogenic to a pathogenic stage may depend on physiological alterations of the host, environmental changes, and/or stress factors. In some woods of the Salento Peninsula (Apulia Region, Italy), sudden declines of holm oak plants (Quercus ilex L.) have been observed since 2016. The morphological and molecular characterization of representative fungal isolates associated with cankers and necrosis in declining plants indicated that these isolates belong to the Botryosphaeriaceae family, and the most frequent species were Diplodia corticola and Diplodia quercivora, followed by Neofusicoccum vitifusiforme. In artificially inoculated young holm oak plants, both D. corticola and D. quercivora species produced intense and severe subcortical and leaf margin necrosis. N. vitifusiforme, although less aggressive, induced the same symptoms. Our research, in addition to confirming the involvement of D. corticola in olm oak decline, represents the first report of D. quercivora as a new pathogen of Q. ilex in Italy. Furthermore, to the best of our knowledge, we also found N. vitifusiforme as a new pathogen of Q. ilex.

Impact of an Extremely Dry Period on Tree Defoliation and Tree Mortality in Serbia.

This paper presents research results on forest decline in Serbia. The results were obtained through monitoring defoliation of 34 tree species at 130 sample plots during the period from 2004 to 2018. This research aimed to determine whether the occurrence of defoliation and tree mortality were caused by drought. Defoliation was assessed in 5% steps according to the International Co-operative Programme on Assessment and Monitoring of Air Pollution Effects on Forests (ICP Forests) methodology. All the trees recorded as dead were singled out, and annual mortality rates were calculated. To determine changes in air temperature and precipitation regimes during the study period, we processed and analysed climatic data related to air temperature and precipitation throughout the year and in the growing season at 28 main weather stations in Serbia. Tree mortality patterns were established by classifying trees into three groups. The first group of trees exhibited a gradual increase in defoliation during the last few years of monitoring, with dying as the final outcome. The second group was characterised by sudden death of trees. The third group of trees reached a higher degree of defoliation immediately after the first monitoring year, and the trees died after several years. Tree mortality rates were compared between years using the Standardised Precipitation Evaporation Index (SPI) and the Standardised Precipitation Evapotranspiration Index (SPEI), the most common methods used to monitor drought. The most intensive forest decline was recorded during the period from 2013 to 2016, when the largest percentage of the total number of all trees died. According to the annual mortality rates calculated for the three observation periods (2004-2008, 2009-2013, and 2014-2018) the highest forest decline rate was recorded in the period from 2014 to 2018, with no statistically significant difference between broadleaved and coniferous tree species. As the sample of coniferous species was small, the number of sample plots should be increased in order to achieve better systematic forest condition monitoring in Serbia. The analysis of the relationship between defoliation and climatic parameters proved the correlation between them. It was noted that the forest decline in Serbia was preceded by an extremely dry period with high temperatures from 2011 to 2013, supporting the hypothesis that it was caused by drought. We therefore conclude that these unfavourable climatic conditions had serious and long-term consequences on forest ecosystems in Serbia.

Post-drought conditions and hydraulic dysfunction determine tree resilience and mortality across Mediterranean Aleppo pine (Pinus halepensis) populations after an extreme drought event.

Drought-related tree mortality is a global phenomenon that currently affects a wide range of forests. Key functional variables on plant hydraulics, carbon economy, growth and allocation have been identified and play a role in tree drought responses. However, tree mortality thresholds based on such variables are difficult to identify, especially under field conditions. We studied several Aleppo pine populations differently affected by an extreme drought event in 2014, with mortality rates ranging from no mortality to 90% in the most severely affected population. We hypothesized that mortality is linked with high levels of xylem embolism, i.e., hydraulic dysfunction, which would also lead to lower tree resistance to drought in subsequent years. Despite not finding any differences among populations in the vulnerability curves to xylem embolism, there were large differences in the hydraulic safety margin (HSM) and the hydraulic dysfunction level. High mortality rates were associated with a negative HSM when xylem embolism reached values over 60%. We also found forest weakening and post-drought mortality related to a low hydraulic water transport capacity, reduced plant growth, low carbohydrate contents and high pest infestation rates. Our results highlight the importance of drought severity and the hydraulic dysfunction level on pine mortality, as well as post-drought conditions during recovery processes.