Long-term soil warming decreases soil microbial necromass carbon by adversely affecting its production and decomposition.

Microbial necromass carbon (MNC) accounts for a large fraction of soil organic carbon (SOC) in terrestrial ecosystems. Yet our understanding of the fate of this large carbon pool under long-term warming is uncertain. Here, we show that 14 years of soil warming (+4°C) in a temperate forest resulted in a reduction in MNC by 11% (0-10 cm) and 33% (10-20 cm). Warming caused a decrease in the content of MNC due to a decline in microbial biomass carbon and reduced microbial carbon use efficiency. This reduction was primarily caused by warming-induced limitations in available soil phosphorus, which, in turn, constrained the production of microbial biomass. Conversely, warming increased the activity of soil extracellular enzymes, specifically N-acetylglucosaminidase and leucine aminopeptidase, which accelerated the decomposition of MNC. These findings collectively demonstrate that decoupling of MNC formation and decomposition underlie the observed MNC loss under climate warming, which could affect SOC content in temperate forest ecosystems more widespread.

Major tree species of Central European forests differ in their proportion of positive, negative, and nonstationary growth trends.

Temperate forests are undergoing significant transformations due to the influence of climate change, including varying responses of different tree species to increasing temperature and drought severity. To comprehensively understand the full range of growth responses, representative datasets spanning extensive site and climatic gradients are essential. This study utilizes tree-ring data from 550 sites from the temperate forests of Czechia to assess growth trends of six dominant Central European tree species (European beech, Norway spruce, Scots pine, silver fir, sessile and pedunculate oak) over 1990-2014. By modeling mean growth series for each species and site, and employing principal component analysis, we identified the predominant growth trends. Over the study period, linear growth trends were evident across most sites (56% increasing, 32% decreasing, and 10% neutral). The proportion of sites with stationary positive trends increased from low toward high elevations, whereas the opposite was true for the stationary negative trends. Notably, within the middle range of their distribution (between 500 and 700 m a.s.l.), Norway spruce and European beech exhibited a mix of positive and negative growth trends. While Scots pine growth trends showed no clear elevation-based pattern, silver fir and oaks displayed consistent positive growth trends regardless of site elevation, indicating resilience to the ongoing warming. We demonstrate divergent growth trajectories across space and among species. These findings are particularly important as recent warming has triggered a gradual shift in the elevation range of optimal growth conditions for most tree species and has also led to a decoupling of growth trends between lowlands and mountain areas. As a result, further future shifts in the elevation range and changes in species diversity of European temperate forests can be expected.

Differences in spatial niche of terrestrial mammals when facing extreme snowfall: the case in east Asian forests.

BACKGROUND: Recent climate changes have produced extreme climate events. This study focused on extreme snowfall and intended to discuss the vulnerability of temperate mammals against it through interspecies comparisons of spatial niches in northern Japan. We constructed niche models for seven non-hibernating species through wide-scaled snow tracking on skis, whose total survey length was 1144 km. RESULTS: We detected a low correlation (rs < 0.4) between most pairs of species niches, indicating that most species possessed different overwintering tactics. A morphological advantage in locomotion cost on snow did not always expand niche breadth. In contrast, a spatial niche could respond to (1) drastic landscape change by a diminishing understory due to snow, possibly leading to changes in predator-prey interactions, and (2) the mass of cold air, affecting thermoregulatory cost and food accessibility. When extraordinary snowfall occurred, the nonarboreal species with larger body sizes could niche shift, whereas the smaller-sized or semi-arboreal mammals did not. In addition, compared to omnivores, herbivores were prone to severe restriction of niche breadth due to a reduction in food accessibility under extreme climates. CONCLUSIONS: Dietary habits and body size could determine the redundancy of niche width, which may govern robustness/vulnerability to extreme snowfall events.

Evaluating the definition and distribution of spring ephemeral wildflowers in eastern North America.

PREMISE: The herbaceous layer accounts for the majority of plant biodiversity in eastern North American forests, encompassing substantial variation in life history strategy and function. One group of early-season herbaceous understory species, colloquially referred to as spring ephemeral wildflowers, are ecologically and culturally important, but little is known about the prevalence and biogeographic patterns of the spring ephemeral strategy. METHODS: We used observations collected by the Global Biodiversity Information Facility (GBIF) to quantify the ephemerality of 559 understory forb species across eastern North America and classify them according to a continuous ephemerality index (ranging from 0 = never ephemeral to 1 = always ephemeral). We then used this information to model where ephemeral forbs were most common across the landscape with the goal of identifying geographic and environmental drivers important to their distributions and ranges. RESULTS: Only 3.4% of all understory wildflower species were spring ephemerals in all parts of their range, and 18.4% (103 species) were ephemeral in at least part of their range. Spring ephemerals peaked in absolute species richness and relative proportion at mid latitudes. CONCLUSIONS: Spring ephemeral phenology is an important shade-avoidance strategy for a large segment of the total understory species in temperate deciduous forests. In North America, the strategy is relatively most important for forest understories at mid latitudes. The definitions of spring ephemerality we provide here serve as an important ecological context for conservation priorities and to evaluate responses of this biodiverse group to future environmental change.

Morphological and molecular data on the bat flies, Basilia silvae (Brèthes, 1913) and Trichobius parasiticus Gervais, 1844 parasitizing Chilean bats.

Among mammals, bats harbour the greatest taxonomic diversity of ectoparasitic arthropods. This is mainly due to their high mobility, wide distribution range and gregarious social behaviour. In Chile, 17 species of bats have been reported; however, their ectoparasitic arthropofauna has been little studied. There are currently 12 taxa recorded, belonging to two classes and associated with only four species of bats. This study aimed to investigate the diversity of parasitic flies associated with bats in three ecoregions: Atacama Desert, Chilean Matorral and Valdivian temperate forest. During 2021, using mist nets, bats were captured in Anzota caves (Atacama Desert ecoregion), Huelquén and Alto Jahuel (Chilean Matorral ecoregion), Cherquenco and San Patricio (Valdivian temperate forest). Each bat was carefully checked during a 15-min interval for the collection of ectoparasites. The arthropods were deposited in vials with 96% ethanol. A total of 26 bats corresponding to three species (Vespertilionidae: Histiotus magellanicus Philippi, 1866, Myotis arescens (Osgood, 1943); Phyllostomidae: Desmodus rotundus (Geoffroy, 1810)) were captured from which a total of 142 ectoparasitic arthropods were collected. Bat flies were separated/identified under a stereomicroscope. Additionally, from the fieldwork, we report the presence of other ectoparasites associated with Chilean bats. In our study, we report new host-parasite associations between Trichobius parasiticus Gervais, 1844 (Diptera: Streblidae) on D. rotundus, and Basilia silvae (Brèthes, 1913) (Diptera: Nycteribiidae) in M. arescens in Chile. Our study extended the latitudinal range of distribution for B. silvae to Araucanía region, and we report for first time T. parasiticus in the country. Additionally, partial sequences of the cytochrome c oxidase I gene were obtained from these specimens. Although there is slight morphological variation in the specimens of T. parasiticus, phylogenetic analyses suggest that they correspond to the same species. The sequences generated for B. silvae represent the first for the species. Authors recommend the use of an integrative approach in the identification of ectoparasites in poorly studied ecoregions and hosts. The integration of different markers is necessary to determine more precisely the phylogenetic relationships between South American populations and species of the genera Basilia and Trichobius.

Altitudinal Impact on Phytochemical Composition and Mycorrhizal Diversity of Taxus Contorta Griff in the Temperate Forest of Shimla District.

Taxus contorta (family Taxaceae) is a native plant of temperate region of western Himalaya. The current study investigated the effect of altitude on the phytochemical composition and mycorrhizal diversity, associated with distribution of T. contorta in Shimla district, Himachal Pradesh, India. Quantitative phytochemical analysis of the leaf extracts indicated that alkaloid levels decreased with altitude, with the highest value in Himri's methanol extracts (72.79 ± 1.08 mg/g) while phenol content increased with altitude, peaking in Nankhari's methanol extracts (118.83 ± 5.90 mg/g). Saponin content was higher in methanol extracts (78.13 ± 1.66 mg/g in Nankhari, 68.06 ± 1.92 mg/g in Pabbas, and 56.32 ± 1.93 mg/g in Himri). Flavonoid levels were notably higher in chloroform extracts, particularly in Nankhari (219.97 ± 2.99 mg/g), and positively correlated with altitude. Terpenoids were higher in chloroform extracts at Himri (11.34 ± 0.10 mg/g) and decreased with altitude. Taxol content showed minimal variation between solvents and altitudes (4.53-6.98 ppm), while rutin was only detected in methanol extracts (1.31-1.46 ppm). Mycorrhizal spore counts in T. contorta's rhizosphere varied with altitude: highest at Himri (77.83 ± 2.20 spores/50 g soil), decreasing to Pabbas (68.06 ± 1.96 spores/50 g soil) and lowest at Nankhari (66.00 ± 2.77 spores/50 g soil), with 17 AMF species identified overall, showing significant altitudinal influence on spore density. The rhizosphere of T. contorta was shown to be dominated by the Glomus species. The rhizospheric soil of the plant was found to be slightly acidic. Organic carbon and available potassium content decreased contrasting with increasing available nitrogen and phosphorus with altitude. Correlation data showed strong negative links between organic carbon (-0.83), moderate positive for nitrogen (0.46) and phosphorus (0.414), and moderate negative for potassium (-0.56) with the altitude. This study provides a comprehensive insight into changes in phytochemical constituents, mycorrhizal diversity and soil composition of T. contorta along a range of altitude.

Are absorptive root traits good predictors of ecosystem functioning? A test in a natural temperate forest.

Trait-based approaches provide a useful framework to predict ecosystem functions under intensifying global change. However, our current understanding of trait-functioning relationships mainly relies on aboveground traits. Belowground traits (e.g. absorptive root traits) are rarely studied although these traits are related to important plant functions. We analyzed four pairs of analogous leaf and absorptive root traits of woody plants in a temperate forest and examined how these traits are coordinated at the community-level, and to what extent the trait covariation depends on local-scale environmental conditions. We then quantified the contributions of leaf and absorptive root traits and the environmental conditions in determining two important forest ecosystem functions, aboveground carbon storage, and woody biomass productivity. The results showed that both morphological trait pairs and chemical trait pairs exhibited positive correlations at the community level. Absorptive root traits show a strong response to environmental conditions compared to leaf traits. We also found that absorptive root traits were better predictors of the two forest ecosystem functions than leaf traits and environmental conditions. Our study confirms the important role of belowground traits in modulating ecosystem functions and deepens our understanding of belowground responses to changing environmental conditions.

Differences between tree stem CO2 efflux and O2 influx rates cannot be explained by internal CO2 transport or storage in large beech trees.

Tree stem respiration (RS ) is a substantial component of the forest carbon balance. The mass balance approach uses stem CO2 efflux and internal xylem fluxes to sum up RS , while the oxygen-based method assumes O2 influx as a proxy of RS . So far, both approaches have yielded inconsistent results regarding the fate of respired CO2 in tree stems, a major challenge for quantifying forest carbon dynamics. We collected a data set of CO2 efflux, O2 influx, xylem CO2 concentration, sap flow, sap pH, stem temperature, nonstructural carbohydrates concentration and potential phosphoenolpyruvate carboxylase (PEPC) capacity on mature beech trees to identify the sources of differences between approaches. The ratio of CO2 efflux to O2 influx was consistently below unity (0.7) along a 3-m vertical gradient, but internal fluxes did not bridge the gap between influx and efflux, nor did we find evidence for changes in respiratory substrate use. PEPC capacity was comparable with that previously reported in green current-year twigs. Although we could not reconcile differences between approaches, results shed light on the uncertain fate of CO2 respired by parenchyma cells across the sapwood. Unexpected high values of PEPC capacity highlight its potential relevance as a mechanism of local CO2 removal, which merits further research.

Urbanization and fragmentation have opposing effects on soil nitrogen availability in temperate forest ecosystems.

Nitrogen (N) availability relative to plant demand has been declining in recent years in terrestrial ecosystems throughout the world, a phenomenon known as N oligotrophication. The temperate forests of the northeastern U.S. have experienced a particularly steep decline in bioavailable N, which is expected to be exacerbated by climate change. This region has also experienced rapid urban expansion in recent decades that leads to forest fragmentation, and it is unknown whether and how these changes affect N availability and uptake by forest trees. Many studies have examined the impact of either urbanization or forest fragmentation on nitrogen (N) cycling, but none to our knowledge have focused on the combined effects of these co-occurring environmental changes. We examined the effects of urbanization and fragmentation on oak-dominated (Quercus spp.) forests along an urban to rural gradient from Boston to central Massachusetts (MA). At eight study sites along the urbanization gradient, plant and soil measurements were made along a 90 m transect from a developed edge to an intact forest interior. Rates of net ammonification, net mineralization, and foliar N concentrations were significantly higher in urban than rural sites, while net nitrification and foliar C:N were not different between urban and rural forests. At urban sites, foliar N and net ammonification and mineralization were higher at forest interiors compared to edges, while net nitrification and foliar C:N were higher at rural forest edges than interiors. These results indicate that urban forests in the northeastern U.S. have greater soil N availability and N uptake by trees compared to rural forests, counteracting the trend for widespread N oligotrophication in temperate forests around the globe. Such increases in available N are diminished at forest edges, however, demonstrating that forest fragmentation has the opposite effect of urbanization on coupled N availability and demand by trees.

Sources, characteristics, and in situ degradation of dissolved organic matters: A case study of a drinking water reservoir located in a cold-temperate forest.

Dissolved organic matter (DOM) plays a pivotal role in the biogeochemical cycles of elements and the regulation of forest ecosystem functions. However, studies on the regional and seasonal characteristics of DOM in cold-temperate montane forests are still not comprehensive. In this study, samples of water, soil, and sediment from different sites in the forest drainage basin were collected, and their DOM was characterized by an excitation-emission matrix and parallel factor analysis (EEM-PARAFAC). The results showed that terrestrial-sourced humic-like substances were the dominant DOM in the studied reservoir and inflowing rivers. The quality and quantity of DOM exhibited spatiotemporal variations with the influence of terrain and monsoonal precipitation. The average concentration of dissolved organic carbon (DOC) in the wet season was 11.62 mg/L, which was higher than that in the dry season (8.18 mg/L). Higher humification index (HIX) values were observed in the wet season and upstream water than in the dry season and reservoir water. Fourier transform ion cyclotron resonance mass spectrometry (FTICR-MS) was used to further develop a molecular-level understanding of the in situ degradation process of DOM. The results indicated that photodegradation rather than biodegradation may play a dominant role in the in situ degradation of terrestrial-sourced humic-like substances under natural conditions. The biodegradability of DOM was enhanced after the in situ degradation process. Additionally, a significant decrease in the precursors of disinfectant byproducts in DOM was observed after in situ degradation. To our knowledge, this is the first study of the sources, characteristics, and in situ degradation of DOM in a reservoir in a cold-temperate forest. These findings help better understand the quality, quantity, and biogeochemical process of DOM in the studied reservoir and may contribute to the selection of drinking water treatment technologies for water supply.

Phenological responses of temperate and boreal trees to warming depend on ambient spring temperatures, leaf habit, and geographic range.

Changes in plant phenology associated with climate change have been observed globally. What is poorly known is whether and how phenological responses to climate warming will differ from year to year, season to season, habitat to habitat, or species to species. Here, we present 5 y of phenological responses to experimental warming for 10 subboreal tree species. Research took place in the open-air B4WarmED experiment in Minnesota. The design is a two habitat (understory and open) × three warming treatments (ambient, +1.7 °C, +3.4 °C) factorial at two sites. Phenology was measured twice weekly during the growing seasons of 2009 through 2013. We found significant interannual variation in the effect of warming and differences among species in response to warming that relate to geographic origin and plant functional group. Moreover, responses to experimental temperature variation were similar to responses to natural temperature variation. Warming advanced the date of budburst more in early compared to late springs, suggesting that to simulate interannual variability in climate sensitivity of phenology, models should employ process-based or continuous development approaches. Differences among species in timing of budburst were also greater in early compared to late springs. Our results suggest that climate change-which will make most springs relatively "early"-could lead to a future with more variable phenology among years and among species, with consequences including greater risk of inappropriately early leafing and altered interactions among species.

The effects of intrinsic water-use efficiency and climate on wood anatomy.

Climate warming may induce growth decline in warm-temperate areas subjected to seasonal soil moisture deficit, whereas increasing atmospheric CO2 concentration (Ca) is expected to enhance tree growth. An accurate understanding of tree growth and physiological processes responding to climate warming and increasing Ca is critical. Here, we analyzed tree-ring stable carbon isotope and wood anatomical traits of Pinus tabuliformis from Qinling Mountains in China to understand how lumen diameter (LD) determining potential hydraulic conductivity and cell-wall thickness (CWT) determining carbon storage responded to climate and Ca. The effects of climate and Ca on intrinsic water-use efficiency (iWUE) were isolated, and iWUE values due to only-climate (iWUEClim) and only-CO2 effects (iWUECO2) were obtained. During a low-iWUE period, the influences of climate on earlywood (EW) LD and latewood (LW) CWT prevailed. During a high-iWUE period, CO2 fertilization promoted cell enlargement and carbon storage but this was counteracted by a negative influence of climate warming. The limiting direct effects of iWUEClim and indirect effects of climate on EW LD were greater than on LW CWT. P. tabuliformis in temperate forests will face a decline of growth and carbon fixation, but will produce embolism-resistant tracheids with narrow lumen responding to future hotter droughts.

Effect of particle sizes of biochar on CO2 emissions in a poplar plantation of ancient Yellow River channel, China.

Forest soil is a vital pool of organic carbon, which is sensitive to management. Biochar addition could change the CO2 emissions from soil, but its effects are still ambiguous. Moreover, the impacts of particle sizes of biochar on CO2 emissions are still unknown. In this study, a series of field experiments were conducted to investigate the effects of biochar addition on CO2 emissions in a poplar plantation (Populus nigra), China. Biochar with two application rates of (10 and 50 t/ha) and three particle sizes (3-1 mm, 1-0.1 mm, and <0.1 mm) was applied into the surface soil (0-10 cm), and the soil without biochar was set as control. The results showed that a high level of fine biochar addition (1-0.1 mm and <0.1 mm) had similar and positive effects on CO2 emissions by increasing the contents of soil ammonium, available phosphorus, easily oxidizable carbon, soil moisture, soil capillary pore, and the activity of ß-glucosidase. However, biochar addition (1-0.1 mm and <0.1 mm) reduced the bioavailability of dissolved organic carbon (DOC), producing a negative relationship between DOC content and CO2 emissions. This investigation highlights the importance of biochar with different particle sizes in adjusting CO2 emissions from temperate soils.

Carbon sink and source function of Eastern Himalayan forests: implications of change in climate and biotic variables.

Forests serve as a sink and source of carbon and play a substantial role in regional and global carbon cycling. The Himalayan forests act as climate regulators of the Hindukush region, which is experiencing climate change at a high pace, and a proper understanding of these systems is necessary to mitigate this problem. We hypothesize that the variance of abiotic factors and vegetation will influence the carbon sink and source function of the different forest types of the Himalayas. Carbon sequestration was computed from the increment of carbon stocks estimated allometrically using Forest Survey of India equations, and soil CO2 flux was determined by the alkali absorption method. The carbon sequestration rate and CO2 flux by the different forests exhibited a negative relation. The carbon sequestration rate was highest with minimum emission in the temperate forest, while the tropical forest recorded the least sequestration and maximum carbon flux rate. The Pearson correlation test between carbon sequestration and tree species richness and diversity revealed a positive-significant influence but negative relation with climatic factors. An analysis of variance indicated significant seasonal differences between the rate of soil carbon emissions due to variations in the forest. A multivariate regression analysis of the monthly soil CO2 emission rate shows high variability (85%) due to fluctuations of climatic variables in the Eastern Himalayan forests. Results of the present study revealed that the carbon sink and source function of forests respond to changes in forest types, climatic variables, and edaphic factors. Tree species and soil nutrient content influenced carbon sequestration, while shifts in climatic factors influenced soil CO2 emission rate. Increased temperature and rainfall may further change the soil quality by enhancing soil CO2 emission and reducing soil organic carbon, thereby impacting this region's carbon sink and source function. Enhancing tree diversity in the forests of this region may be beneficial for retarding this impact.

Limitation by vapour pressure deficit shapes different intra-annual growth patterns of diffuse- and ring-porous temperate broadleaves.

Understanding the effects of temperature and moisture on radial growth is vital for assessing the impacts of climate change on carbon and water cycles. However, studies observing growth at sub-daily temporal scales remain scarce. We analysed sub-daily growth dynamics and its climatic drivers recorded by point dendrometers for 35 trees of three temperate broadleaved species during the years 2015-2020. We isolated irreversible growth driven by cambial activity from the dendrometer records. Next, we compared the intra-annual growth patterns among species and delimited their climatic optima. The growth of all species peaked at air temperatures between 12 and 16°C and vapour pressure deficit (VPD) below 0.1 kPa. Acer pseudoplatanus and Fagus sylvatica, both diffuse-porous, sustained growth under suboptimal VPD. Ring-porous Quercus robur experienced a steep decline of growth rates with reduced air humidity. This resulted in multiple irregular growth peaks of Q. robur during the year. By contrast, the growth patterns of the diffuse-porous species were always right-skewed unimodal with a peak in June between day of the year 150-170. Intra-annual growth patterns are shaped more by VPD than temperature. The different sensitivity of radial growth to VPD is responsible for unimodal growth patterns in both diffuse-porous species and multimodal growth pattern in Q. robur.

Cutting tree rings into time slices: how intra-annual dynamics of wood formation help decipher the space-for-time conversion.

Tree-ring anatomy, microdensity and isotope records provide valuable intra-annual information. However, extracting signals at that scale is challenged by the complexity of xylogenesis, where two major processes - cell enlargement and wall thickening - occur at different times and rates. We characterized the space-for-time association in the tree rings of three conifer species by examining the duration, overlapping, inter-tree synchronicity and interannual stability during cell enlargement and wall thickening across regular tree-ring sectors (portions of equal tangential width). The number of cells and cell differentiation rates determined the duration of sector formation, which augmented more rapidly throughout the ring for wall thickening than for enlargement. Increasing the number of sectors above c. 15 had a limited effect on improving time resolution because consecutive sector formation overlapped greatly in time, especially in narrow rings and during wall thickening. Increasing the number of sectors also resulted in lower synchronicity and stability of intermediate-sector enlargement, whereas all sectors showed high synchronicity and stability during wall thickening. Increasing the number of sectors had a stronger effect on enhancing time-series resolution for enlargement- than for wall-thickening-related traits, which would nevertheless produce more reliable intra-annual chronologies as a result of the more similar calendars across trees and years in wall thickening.

Inter-annual and inter-species tree growth explained by phenology of xylogenesis.

Wood formation determines major long-term carbon (C) accumulation in trees and therefore provides a crucial ecosystem service in mitigating climate change. Nevertheless, we lack understanding of how species with contrasting wood anatomical types differ with respect to phenology and environmental controls on wood formation. In this study, we investigated the seasonality and rates of radial growth and their relationships with climatic factors, and the seasonal variations of stem nonstructural carbohydrates (NSC) in three species with contrasting wood anatomical types (red oak: ring-porous; red maple: diffuse-porous; white pine: coniferous) in a temperate mixed forest during 2017-2019. We found that the high ring width variability observed in both red oak and red maple was caused more by changes in growth duration than growth rate. Seasonal radial growth patterns did not vary following transient environmental factors for all three species. Both angiosperm species showed higher concentrations and lower inter-annual fluctuations of NSC than the coniferous species. Inter-annual variability of ring width varied by species with contrasting wood anatomical types. Due to the high dependence of annual ring width on growth duration, our study highlights the critical importance of xylem formation phenology for understanding and modelling the dynamics of wood formation.

Carbon allocation to root exudates is maintained in mature temperate tree species under drought.

Carbon (C) exuded via roots is proposed to increase under drought and facilitate important ecosystem functions. However, it is unknown how exudate quantities relate to the total C budget of a drought-stressed tree, that is, how much of net-C assimilation is allocated to exudation at the tree level. We calculated the proportion of daily C assimilation allocated to root exudation during early summer by collecting root exudates from mature Fagus sylvatica and Picea abies exposed to experimental drought, and combining above- and belowground C fluxes with leaf, stem and fine-root surface area. Exudation from individual roots increased exponentially with decreasing soil moisture, with the highest increase at the wilting point. Despite c. 50% reduced C assimilation under drought, exudation from fine-root systems was maintained and trees exuded 1.0% (F. sylvatica) to 2.5% (P. abies) of net C into the rhizosphere, increasing the proportion of C allocation to exudates two- to three-fold. Water-limited P. abies released two-thirds of its exudate C into the surface soil, whereas in droughted F. sylvatica it was only one-third. Across the entire root system, droughted trees maintained exudation similar to controls, suggesting drought-imposed belowground C investment, which could be beneficial for ecosystem resilience.

Regeneration from seed in herbaceous understorey of ancient woodlands of temperate Europe.

BACKGROUND AND AIMS: European ancient woodlands are subject to land use change, and the distribution of herbaceous understorey species may be threatened because of their poor ability to colonize isolated forest patches. The regeneration niche can determine the species assembly of a community, and seed germination traits may be important descriptors of this niche. METHODS: We analysed ecological records for 208 herbaceous species regarded as indicators of ancient woodlands in Europe and, where possible, collated data on seed germination traits, reviewed plant regeneration strategies and measured seed internal morphology traits. The relationship between plant regeneration strategies and ecological requirements was explored for 57 species using ordination and classification analysis. KEY RESULTS: Three regeneration strategies were identified. Species growing in closed-canopy areas tend to have morphological seed dormancy, often requiring darkness and low temperatures for germination, and their shoots emerge in early spring, thus avoiding the competition for light from canopy species. These species are separated into two groups: autumn and late winter germinators. The third strategy is defined by open-forest plants with a preference for gaps, forest edges and riparian forests. They tend to have physiological seed dormancy and germinate in light and at higher temperatures, so their seedlings emerge in spring or summer. CONCLUSION: Seed germination traits are fundamental to which species are good or poor colonizers of the temperate forest understorey and could provide a finer explanation than adult plant traits of species distribution patterns. Seed dormancy type, temperature stratification and light requirements for seed germination are important drivers of forest floor colonization patterns and should be taken in account when planning successful ecological recovery of temperate woodland understories.

Empirically validated drought vulnerability mapping in the mixed conifer forests of the Sierra Nevada.

Severe droughts are predicted to become more frequent in the future, and the consequences of such droughts on forests can be dramatic, resulting in massive tree mortality, rapid change in forest structure and composition, and substantially increased risk of catastrophic fire. Forest managers have tools at their disposal to try to mitigate these effects but are often faced with limited resources, forcing them to make choices about which parts of the landscape to target for treatment. Such planning can greatly benefit from landscape vulnerability assessments, but many existing vulnerability analyses are unvalidated and not grounded in robust empirical datasets. We combined robust sets of ground-based plot and remote sensing data, collected during the 2012-2016 California drought, to develop rigorously validated tools for assessing forest vulnerability to drought-related canopy tree mortality for the mixed conifer forests of the Sequoia and Kings Canyon national parks and potentially for mixed conifer forests in the Sierra Nevada as a whole. Validation was carried out using a large external dataset. The best models included normalized difference vegetation index (NDVI), elevation, and species identity. Models indicated that tree survival probability decreased with greenness (as measured by NDVI) and elevation, particularly if trees were growing slowly. Overall, models showed good calibration and validation, especially for Abies concolor, which comprise a large majority of the trees in many mixed conifer forests in the Sierra Nevada. Our models tended to overestimate mortality risk for Calocedrus decurrens and underestimate risk for pine species, in the latter case probably due to pine bark beetle outbreak dynamics. Validation results indicated dangers of overfitting, as well as showing that the inclusion of trees already under attack by bark beetles at the time of sampling can give false confidence in model strength, while also biasing predictions. These vulnerability tools should be useful to forest managers trying to assess which parts of their landscape were vulnerable during the 2012-2016 drought, and, with additional validation, may prove useful for ongoing assessments and predictions of future forest vulnerability.