A genome-guided strategy for climate resilience in American chestnut restoration populations.

American chestnut (Castanea dentata) is a deciduous tree species of eastern North America that was decimated by the introduction of the chestnut blight fungus (Cryphonectria parasitica) in the early 20th century. Although millions of American chestnuts survive as root collar sprouts, these trees rarely reproduce. Thus, the species is considered functionally extinct. American chestnuts with improved blight resistance have been developed through interspecific hybridization followed by conspecific backcrossing, and by genetic engineering. Incorporating adaptive genomic diversity into these backcross families and transgenic lines is important for restoring the species across broad climatic gradients. To develop sampling recommendations for ex situ conservation of wild adaptive genetic variation, we coupled whole-genome resequencing of 384 stump sprouts with genotype-environment association analyses and found that the species range can be subdivided into three seed zones characterized by relatively homogeneous adaptive allele frequencies. We estimated that 21 to 29 trees per seed zone will need to be conserved to capture most extant adaptive diversity. We also resequenced the genomes of 269 backcross trees to understand the extent to which the breeding program has already captured wild adaptive diversity, and to estimate optimal reintroduction sites for specific families on the basis of their adaptive portfolio and future climate projections. Taken together, these results inform the development of an ex situ germplasm conservation and breeding plan to target blight-resistant breeding populations to specific environments and provides a blueprint for developing restoration plans for other imperiled tree species.

Legacy effects of premature defoliation in response to an extreme drought event modulate phytochemical profiles with subtle consequences for leaf herbivory in European beech.

Extreme droughts can have long-lasting effects on forest community dynamics and species interactions. Yet, our understanding of how drought legacy modulates ecological relationships is just unfolding. We tested the hypothesis that leaf chemistry and herbivory show long-term responses to premature defoliation caused by an extreme drought event in European beech (Fagus sylvatica L.). For two consecutive years after the extreme European summer drought in 2018, we collected leaves from the upper and lower canopy of adjacently growing drought-stressed and unstressed trees. Leaf chemistry was analyzed and leaf damage by different herbivore-feeding guilds was quantified. We found that drought had lasting impacts on leaf nutrients and on specialized metabolomic profiles. However, drought did not affect the primary metabolome. Drought-related phytochemical changes affected damage of leaf-chewing herbivores whereas damage caused by other herbivore-feeding guilds was largely unaffected. Drought legacy effects on phytochemistry and herbivory were often weaker than between-year or between-canopy strata variability. Our findings suggest that a single extreme drought event bears the potential to long-lastingly affect tree-herbivore interactions. Drought legacy effects likely become more important in modulating tree-herbivore interactions since drought frequency and severity are projected to globally increase in the coming decades.

The interacting effect of climate change and herbivory can trigger large-scale transformations of European temperate forests.

In many regions of Europe, large wild herbivores alter forest community composition through their foraging preferences, hinder the forest's natural adaptive responses to climate change, and reduce ecosystem resilience. We investigated a widespread European forest type, a mixed forest dominated by Picea abies, which has recently experienced an unprecedented level of disturbance across the continent. Using the forest landscape model iLand, we investigated the combined effect of climate change and herbivory on forest structure, composition, and carbon and identified conditions leading to ecosystem transitions on a 300-year timescale. Eight climate change scenarios, driven by Representative Concentration Pathways 4.5 and 8.5, combined with three levels of regeneration browsing, were tested. We found that the persistence of the current level of browsing pressure impedes adaptive changes in community composition and sustains the presence of the vulnerable yet less palatable P. abies. These development trajectories were tortuous, characterized by a high disturbance intensity. On the contrary, reduced herbivory initiated a transformation towards the naturally dominant broadleaved species that was associated with an increased forest carbon and a considerably reduced disturbance. The conditions of RCP4.5 combined with high and moderate browsing levels preserved the forest within its reference range of variability, defining the actual boundaries of resilience. The remaining combinations of browsing and climate change led to ecosystem transitions. Under RCP4.5 with browsing effects excluded, the new equilibrium conditions were achieved within 120 years, whereas the stabilization was delayed by 50-100 years under RCP8.5 with higher browsing intensities. We conclude that forests dominated by P. abies are prone to transitions driven by climate change. However, reducing herbivory can set the forest on a stable and predictable trajectory, whereas sustaining the current browsing levels can lead to heightened disturbance activity, extended transition times, and high variability in the target conditions.

Biogeographic context is related to local scale tree demography, co-occurrence and functional differentiation.

Identifying the drivers of community structure and dynamics is a major pursuit in ecology. Emphasis is typically placed on the importance of local scale interactions when attempting to explain these fundamental ecological patterns. However, regional scale phenomena are also important predictors. The importance of regional scale context should be more evident in assemblages where multiple species are close to their range margins. Here, we test the importance of regional scale context using data from a temperate forest plot that contains two species groups - one near its northern range limit and one near its southern range limit. We show the proximity of species to their southern or northern range margins is linked to local scale co-occurrence, similarity in gene expression responses to a key environmental driver, demographic performance and inter-specific variation in conspecific negative density dependence. In sum, many of the key local scale patterns and processes of interest to community ecologists are linked to biogeographic context that is frequently ignored.

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.

Carbon stress causes earlier budbreak in shade-tolerant species and delays it in shade-intolerant species.

PREMISE: Climate change may lead to C stress (negative C balance) in trees. Because nonstructural carbohydrates (NSC) are required during metabolic reactivation in the spring, C stress might delay budbreak timing. This effect is expected to be greater in shade-intolerant than in shade-tolerant species, owing to the faster C economy in the shade-intolerant. METHODS: We experimentally induced C stress in saplings of six temperate tree species that differed in their light requirements by exposing them to either full light or shade from summer to spring, then recorded the date of first budbreak for the individuals. Because the levels of C reserves that represent effective C stress may differ among species, we estimated the degree of C stress by recording survival during the experiment and measuring whole-sapling NSC concentrations after budbreak. RESULTS: Shade reduced NSC concentrations and increased the sugar fraction in the NSC in all species. In the shade, shade-intolerant species had higher mortality and generally lower NSC concentrations than the shade-tolerant species, indicating a trend for more severe C stress in species with faster C economy. In shade-intolerant species, budbreak started earlier and proceeded faster in full light than in shade, but in shade-tolerant species budbreak was delayed in full light. The effects of the light environments on budbreak were not greater in shade-intolerant than in shade-tolerant species. CONCLUSIONS: Our study reveals a correspondence between budbreak responses to light and the light requirements of the species. This finding confirms that C metabolism has a significant role in triggering budbreak and demonstrates that whether C stress accelerates or delays budbreak depends on the species' light requirements.

Evolutionary origins of the eastern North American-Mesoamerican floristic disjunction: Current status and future prospects.

Biogeographic disjunction patterns, where multiple taxa are shared between isolated geographic areas, represent excellent systems for investigating the historical assembly of modern biotas and fundamental biological processes such as speciation, diversification, niche evolution, and evolutionary responses to climate change. Studies on plant genera disjunct across the northern hemisphere, particularly between eastern North America (ENA) and eastern Asia (EAS), have yielded tremendous insight on the geologic history and assembly of rich temperate floras. However, one of the most prevalent disjunction patterns involving ENA forests has been largely overlooked: that of taxa disjunct between ENA and cloud forests of Mesoamerica (MAM), with examples including Acer saccharum, Liquidambar styraciflua, Cercis canadensis, Fagus grandifolia, and Epifagus virginiana. Despite the remarkable nature of this disjunction pattern, which has been recognized for over 75 years, there have been few recent efforts to empirically examine its evolutionary and ecological origins. Here I synthesize previous systematic, paleobotanical, phylogenetic, and phylogeographic studies to establish what is known about this disjunction pattern to provide a roadmap for future research. I argue that this disjunction pattern, and the evolution and fossil record of the Mexican flora more broadly, represents a key missing piece in the broader puzzle of northern hemisphere biogeography. I also suggest that the ENA-MAM disjunction represents an excellent system for examining fundamental questions about how traits and life history strategies mediate plant evolutionary responses to climate change and for predicting how broadleaf temperate forests will respond to the ongoing climatic pressures of the Anthropocene.

The Western Himalayan fir tree ring record of soil moisture in Pakistan since 1855.

Data on historical soil moisture is crucial for assessing and responding to droughts that commonly occur in climate change-affected countries. The Himalayan temperate forests in Pakistan are particularly at risk of climate change. Developing nations lack the means to gather surface soil moisture (SSM) information. Tree rings are one way to bridge this gap. Here, we employed dendrochronological methods on climate-sensitive tree rings from Abies pindrow to reconstruct the SSM in the Western Himalayan mountain region of Pakistan from 1855 to 2020. December (r = 0.41), May (r = 0.40), and June (r = 0.65) SSMs were found to be the limiting factors for A. pindrow growth. However, only the June SSM showed reconstruction possibility (coefficient of efficiency = 0.201 and reduction of error = 0.325). Over the studied period, we found 6 years (wet year) when June SSM was above the threshold of 32.04 (mean + 2 δ) and 1 year (dry year) when June SSM was below the threshold of 21.28 (mean - 2 δ). It was revealed that 1921 and 1917 were the driest and wettest SSM of all time, with means of 19.34 and 36.49, respectively. Our study shows that winter soil moisture is critical for the growing season in the context of climate change. Climate change has broad impacts on tree growth in the Western Himalayas. This study will assist various stakeholders in understanding and managing local and regional climate change.

Interplay of moss cover and seasonal variation regulate soil physicochemical properties and net nitrogen mineralization rates in Central Himalayas, India.

Mosses (Class- Bryopsida) are vital to ecosystem dynamics in numerous biomes, although their effects on soil processes are poorly understood. The interplay of moss cover and seasonal variations in soil processes is still unclear in the Indian Central Himalayas. Therefore, we examined the seasonal variations in net nitrogen (N) mineralization rates and several soil properties under two ground covers (with and without moss cover). We used the ex-situ incubation technique to determine N mineralization rates (Rmin) and standard methodology for soil physical and chemical analysis. During the rainy season, the physical properties of the soil and its nutrients, apart from phosphorus, were higher under moss cover. The winter season, however, showed a different pattern, with soil properties exhibiting higher values in soils without moss cover. Ammonium concentrations were higher under moss cover, while nitrate concentrations were higher in soil without moss cover during rainy and winter seasons. The Rmin rates were higher in soil under moss cover, indicating that moss cover promotes N transformation. In contrast, Rmin rates were negative in soil without moss cover, indicating that N immobilization was dominant in N transformation under this ground cover during the rainy season. Our research shows that mosses positively impact the nutrient status and N mineralization rates in various temperate forest types. The seasonal patterns of soil properties are strongly influenced by soil temperature, moisture, and organic carbon. Therefore, we advocate the conservation of mosses and their integration into forest management plans for better ecosystem processes and services in the ecologically fragile Himalayas.

Assessing the Effectiveness of Silvicultural Treatments on Fire Behavior in the Hyrcanian Temperate Forests of Northern Iran.

We implemented a fire modeling approach to evaluate the effectiveness of silvicultural treatments in reducing potential losses to the Hyrcanian temperate forests of northern Iran, in the Siahkal National Forest (57,110 ha). We compared the effectiveness of selection cutting, low thinning, crown thinning, and clear-cutting treatments implemented during the last ten years (n = 241, 9500-ha) on simulated stand-scale and landscape-scale fire behavior. First, we built a set of fuel models for the different treatment prescriptions. We then modeled 10,000 fires at the 30-m resolution, assuming low, moderate, high, very high, and extreme weather scenarios and human-caused ignition patterns. Finally, we implemented a One-way ANOVA test to analyze stand-level and landscape-scale modeling output differences between treated and untreated conditions. The results showed a significant reduction of stand-level fire hazard, where the average conditional flame length and crown fire probability was reduced by about 12 and 22%, respectively. The conifer plantation patches presented the most significant reduction in the crown fire probability (>35%). On the other hand, we found a minor increase in the overall burn probability and fire size at the landscape scale. Stochastic fire modeling captured the complex interactions among terrain, vegetation, ignition locations, and weather conditions in the study area. Our findings highlight fuel treatment efficacy for moderating potential fire risk and restoring fuel profiles in fire-sensitive temperate forests of northern Iran, where the growing persistent droughts and fuel buildup can lead to extreme fires in the near future.

Functional trait diversity and aboveground biomass of herbaceous vegetation in temperate forests of Kashmir Himalaya.

Studying functional trait diversity can provide crucial clues about the adaptive survival strategies of regional plant species pool. Despite large-scale trait datasets available worldwide, the plant trait data from many biodiversity hotpot regions, like the Himalaya is still scarce. In this study, we aimed to investigate the plant functional traits and aboveground biomass of understory herbaceous vegetation in temperate forests of Overa-Aru wildlife sanctuary of Kashmir Himalaya. We also investigate how these functional traits correlate and what is the magnitude of trait-biomass relationship across the herbaceous species pool. For this, we conducted field sampling and measured leaf functional traits and aboveground biomass of 38 plant species in the study region during peak growing season (July-August) in the year 2021. The results revealed a significant interspecific trait variability among the species studied. We observed a high variability in leaf size and type spectra of the species, with nanophyll and simple leaf lamina, respectively, the most common types among the species studied. The correlation analysis revealed that plant height was positively correlated with aboveground biomass. The variation partitioning analysis revealed that the plant height explained the maximum fraction of variation in aboveground biomass, while least by specific leaf area. Overall, the findings from the present study provide useful insights in understanding trait-trait relationship and trait-environment interaction at the regional scale and can also help in recognizing adaptive functional traits of plant species that determine plant survival under the changing climate in this Himalayan region.

Cumulative growth and stress responses to the 2018-2019 drought in a European floodplain forest.

Droughts increasingly threaten the world's forests and their potential to mitigate climate change. In 2018-2019, Central European forests were hit by two consecutive hotter drought years, an unprecedented phenomenon that is likely to occur more frequently with climate change. Here, we examine tree growth and physiological stress responses (increase in carbon isotope composition; Δδ13 C) to this consecutive drought based on tree rings of dominant tree species in a Central European floodplain forest. Tree growth was not reduced for most species in 2018, indicating that water supply in floodplain forests can partly buffer meteorological water deficits. Drought stress responses in 2018 were comparable to former single drought years but the hotter drought in 2018 induced drought legacies in tree growth while former droughts did not. We observed strong decreases in tree growth and increases in Δδ13 C across all tree species in 2019, which are likely driven by the cumulative stress both consecutive hotter droughts exerted. Our results show that consecutive hotter droughts pose a novel threat to forests under climate change, even in forest ecosystems with comparably high levels of water supply.

Crowding, climate, and the case for social distancing among trees.

In an emerging era of megadisturbance, bolstering forest resilience to wildfire, insects, and drought has become a central objective in many western forests. Climate has received considerable attention as a driver of these disturbances, but few studies have examined the complexities of climate-vegetation-disturbance interactions. Current strategies for creating resilient forests often rely on retrospective approaches, seeking to impart resilience by restoring historical conditions to contemporary landscapes, but historical conditions are becoming increasingly unattainable amidst modern bioclimatic conditions. What becomes an appropriate benchmark for resilience when we have novel forests, rapidly changing climate, and unprecedented disturbance regimes? We combined two longitudinal datasets-each representing some of the most comprehensive spatially explicit, annual tree mortality data in existence-in a post-hoc factorial design to examine the nonlinear relationships between fire, climate, forest spatial structure, and bark beetles. We found that while prefire drought elevated mortality risk, advantageous local neighborhoods could offset these effects. Surprisingly, mortality risk (Pm ) was higher in crowded local neighborhoods that burned in wet years (Pm  = 42%) compared with sparse neighborhoods that burned during drought (Pm  = 30%). Risk of beetle attack was also increased by drought, but lower conspecific crowding impeded the otherwise positive interaction between fire and beetle attack. Antecedent fire increased drought-related mortality over short timespans (<7 years) but reduced mortality over longer intervals. These results clarify interacting disturbance dynamics and provide a mechanistic underpinning for forest restoration strategies. Importantly, they demonstrate the potential for managed fire and silvicultural strategies to offset climate effects and bolster resilience to fire, beetles, and drought.

Using historical spy satellite photographs and recent remote sensing data to identify high-conservation-value forests.

High-conservation-value forests (HCVFs) are critically important for biodiversity and ecosystem service provisioning, but they face many threats. Where systematic HCVF inventories are missing, such as in parts of Eastern Europe, these forests remain largely unacknowledged and therefore often unprotected. We devised a novel, transferable approach for detecting HCVFs based on integrating historical spy satellite images, contemporary remote sensing data (Landsat), and information on current potential anthropogenic pressures (e.g., road infrastructure, population density, demand for fire wood, terrain). We applied the method to the Romanian Carpathians, for which we mapped forest continuity (1955-2019), canopy structural complexity, and anthropogenic pressures. We identified 738,000 ha of HCVF. More than half of this area was identified as susceptible to current anthropogenic pressures and lacked formal protection. By providing a framework for broad-scale HCVF monitoring, our approach facilitates integration of HCVF into forest conservation and management. This is urgently needed to achieve the goals of the European Union's Biodiversity Strategy to maintain valuable forest ecosystems.

Uso de Fotografías Históricas de Satélites Espía y Datos Recientes de Telemetría para Identificar Bosques de Alto Valor para la Conservación Resumen Los bosques de alto valor para la conservación (BAVC) tienen una importancia crítica para el suministro de servicios ambientales y biodiversidad pero enfrentan muchas amenazas. En donde hacen falta inventarios sistemáticos de los BAVC, como en partes del este de Europa, estos bosques siguen siendo ignorados y por lo tanto carecen de protección. Diseñamos una estrategia novedosa y transferible para la detección de BAVC con base en la integración de imágenes de satélites espía, datos contemporáneos de telemetría (Landsat) e información sobre las presiones antropogénicas actuales (p. ej.: infraestructura vial, densidad poblacional, demanda de leña, terreno). Aplicamos el método en los Cárpatos rumanos, para los cuales mapeamos la continuidad forestal (1955 - 2019), la complejidad estructural del dosel y las presiones antropogénicas. Identificamos 738,000 ha de BAVC. Más de la mitad de esta área fue identificada como susceptible a las actuales presiones antropogénicas y además carecía de protección formal. Mediante la aportación de un marco de trabajo para el monitoreo a escala amplia de los BAVC, nuestra estrategia facilita la integración de los BAVC dentro de la gestión y conservación de los bosques. Lo último es una necesidad urgente para alcanzar las metas de la Estrategia de Biodiversidad de la Unión Europea para mantener los ecosistemas boscosos valiosos.

Vertical and horizontal light heterogeneity along gradients of secondary succession in cool- and warm-temperate forests.

Aims: Light availability varies drastically in forests, both vertically and horizontally. Vertical light heterogeneity (i.e., patterns of light attenuation from the forest canopy to the floor) may be related to light competition among trees, while horizontal light heterogeneity (i.e., variations in light intensity at a given height within forests) may be associated with light-niche partitioning among tree species. However, light heterogeneity in vertical and horizontal directions and their associations with forest structure are rarely studied to date. Here we report the first comprehensive study to compare the vertical and horizontal light heterogeneity in differently aged forests in two forest types. Location: Twelve forest stands of different ages in cool-temperate forests (consisting of deciduous broad-leaved trees) and five of different ages in warm-temperate forests (evergreen conifer and deciduous broad-leaved trees) in Japan. Methods: We measured vertical light profiles at 1-m intervals from the understorey (1 m above the ground) to the top canopy (12-22 m depending on stands) at 16 locations for each stand (20 m × 20 m). We also measured structural parameters (diameter at breast height, height, and crown dimensions) for all major trees in these stands. Results: Along the secondary successional gradients, the vertical and horizontal light heterogeneity changed in a systematic manner in both forests. The vertical light attenuation rate was steeper in early succession and more gradual in late succession, and the horizontal light heterogeneity was relatively small in early succession and more pronounced in late succession. The vertical light attenuation rate was different between the two forest types; the light intensity dropped more sharply from the canopy surface in the cool-temperate forests due to the crown being vertically shorter and denser (i.e., higher leaf density per unit volume). Conclusion: In early succession, a steeper light attenuation rate is likely related to the strong light competition among co-occurring trees and thus a self-thinning process. In late succession, the high spatial light heterogeneity in forests (i.e., larger horizontal light heterogeneity and gradual light attenuation rate) may allow more species to partition light, and thus may enhance species coexistence and diversity.

Species co-occurrence and management intensity modulate habitat preferences of forest birds.

BACKGROUND: Species co-occurrences can have profound effects on the habitat use of species, and therefore habitat structure alone cannot fully explain observed abundances. To account for this aspect of community organization, we developed multi-species abundance models, incorporating the local effect of co-occurring and potentially associated species, alongside with environmental predictors, linked mainly to forest management intensity. We coupled it with a landscape-scale analysis to further examine the role of management intensity in modifying the habitat preferences in connection with the landscape context. Using empirical data from the Black Forest in southern Germany, we focused on the forest bird assemblage and in particular on the cavity-nesting and canopy-foraging guilds. We included in the analysis species that co-occur and for which evidence suggests association is likely. RESULTS: Our findings show that the local effect of species associations can mitigate the effects of management intensity on forest birds. We also found that bird species express wider habitat preferences in forests under higher management intensity, depending on the landscape context. CONCLUSIONS: We suspect that species associations may facilitate the utilization of a broader range of environmental conditions under intensive forest management, which benefits some species over others. Networks of associations may be a relevant factor in the effectiveness of conservation-oriented forest management.

Mature conifers assimilate nitrate as efficiently as ammonium from soils in four forest plantations.

Conifers are considered to prefer to take up ammonium (NH4+ ) over nitrate (NO3- ). However, this conclusion is mainly based on hydroponic experiments that separate roots from soils. It remains unclear to what extent mature conifers can use nitrate compared to ammonium under field conditions where both roots and soil microbes compete for nitrogen (N). We conducted an in situ whole mature tree nitrogen-15 (15 N) labeling experiment (15 NH4+ vs 15 NO3- ) over 15 d to quantify ammonium and nitrate uptake and assimilation rates in four 40-yr-old monoculture coniferous plantations (Pinus koraiensis, Pinus sylvestris, Picea koraiensis and Larix olgensis, respectively). For the whole tree, 15 NO3- contributed 39% to 90% to total 15 N tracer uptake among four plantations during the study period. At day 3, the 15 NO3- accounted for 77%, 64%, 62% and 59% by Larix olgensis, Pinus koraiensis, Pinus sylvestris and Picea koraiensis, respectively. Our study indicates that mature coniferous trees assimilated nitrate as efficiently as ammonium from soils even at low soil nitrate concentration, in contrast to the results from hydroponic experiments showing that ammonium uptake dominated over nitrate. This implies that mature conifers can adapt to increasing availability of nitrate in soil, for example, under the context of globalization of N deposition and global warming.

Repetitive seasonal drought causes substantial species-specific shifts in fine-root longevity and spatio-temporal production patterns in mature temperate forest trees.

Temperate forest ecosystems are exposed to a higher frequency, duration and severity of drought. To promote forest longevity in a changing climate, we require a better understanding of the long-term impacts of repetitive drought events on fine-root dynamics in mature forests. Using minirhizotron methods, we investigated the effect of seasonal drought on fine-root dynamics in single-species and mixed-species arrangements of Fagus sylvatica (European beech) and Picea abies (Norway spruce) by means of a 4-yr-long throughfall-exclusion experiment. Fine-root production of both species decreased under drought. However, this reduction was not evident for P. abies when grown intermixed with F. sylvatica. Throughfall-exclusion prolonged the lifespan of P. abies roots but did not change the lifespan of F. sylvatica roots, except in 2016. Fagus sylvatica responded to drought by reducing fine-root production at specific depths and during roof closure. This is the first study to examine long-term trends in mature forest fine-root dynamics under repetitive drought events. Species-specific fine-root responses to drought have implications for the rate and depth of root-derived organic matter supply to soil. From a root dynamics perspective, intermixing tree species is not beneficial to all species but dampens drought impacts on the belowground productivity of P. abies.

Adaptive variation and plasticity in non-structural carbohydrate storage in a temperate tree species.

Trees' total amount of non-structural carbohydrate (NSC) stores and the proportion of these stores residing as insoluble starch are vital traits for individuals living in variable environments. However, our understanding of how stores vary in response to environmental stress is poorly understood as the genetic component of storage is rarely accounted for in studies. Here, we quantified variation in NSC traits in branch samples taken from over 600 clonally transplanted black cottonwood (Populus trichocarpa) trees grown in two common gardens. We found heritable variation in both total NSC stores and the proportion of stores in starch (H2TNC  = 0.19, H2PropStarch  = 0.31), indicating a substantial genetic component of variation. In addition, we found high amounts of plasticity in both traits in response to cold temperatures and significant genotype-by-environment (GxE) interactions in the total amount of NSC stored (54% of P is GxE). This finding of high GxE indicates extensive variation across trees in their response to environment, which may explain why previous studies of carbohydrate stores' responses to stress have failed to converge on a consistent pattern. Overall, we found high amounts of environmental and genetic variation in NSC storage concentrations, which may bolster species against future climate change.

Disentangling the relative importance of spatio-temporal parameters and host specificity in shaping arbuscular mycorrhizal fungus communities in a temperate forest.

Many woody and herbaceous plants in temperate forests cannot establish and survive in the absence of mycorrhizal associations. Most temperate forests are dominated by ectomycorrhizal woody plant species, which implies that the carrying capacity of the habitat for arbuscular mycorrhizal fungi (AMF) is relatively low and AMF could in some cases experience a limitation of propagules. Here we address how the AMF community composition varied in a small temperate forest site in Germany in relation to time, space, two plant host species, and also with regard to the degree to which plots were covered with AMF-associating woody species. The AMF communities in our study were non-random. We observed that space had a greater impact on fungal community composition than either time, mycorrhizal state of the close-by woody species, or the identity of the host plant. The identity of the host plant was the only parameter that modified AMF richness in the roots. The set of parameters which we addressed has rarely been studied together, and the resulting ranking could ease prioritizing some of them to be included in future surveys. AMF are crucial for the establishment of understory plants in temperate forests, making it desirable to further explore how they vary in time and space.