Unpunctual in diversity: The effect of stand species richness on spring phenology of deciduous tree stands varies among species and years.

Climatic drivers alone do not adequately explain the regional variation in budburst timing in deciduous forests across Europe. Stand-level factors, such as tree species richness, might affect budburst timing by creating different microclimates under the same site macroclimate. We assessed different phases of the spring phenology (start, midpoint, end, and overall duration of the budburst period) of four important European tree species (Betula pendula, Fagus sylvatica, Quercus robur and Tilia cordata) in monocultures and four-species mixture stands of a common garden tree biodiversity experiment in Belgium (FORBIO) in 2021 and 2022. Microclimatic differences between the stands in terms of bud chilling, temperature forcing, and soil temperature were considerable, with four-species mixtures being generally colder than monocultures in spring, but not in winter. In the colder spring of 2021, at the stand level, the end of the budburst period was advanced, and its overall duration shortened, in the four-species mixtures. At species level, this response was significant for F. sylvatica. In the warmer spring of 2022, advances in spring phenology in four-species stands were observed again in F. sylvatica and, less markedly, in B. pendula but without a general response at the stand level. Q. robur showed specific patterns with delayed budburst start in 2021 in the four-species mixtures and very short budburst duration for all stands in 2022. Phenological differences between monocultures and four-species mixtures were linked to microclimatic differences in light availability rather than in temperature as even comparatively colder microclimates showed an advanced phenology. Compared to weather conditions, tree species richness had a lower impact on budburst timing, but this impact can be of importance for key species like F. sylvatica and colder springs. These results indicate that forest biodiversity can affect budburst phenology, with wider implications, especially for forest- and land surface models.

Late autumn warming can both delay and advance spring budburst through contrasting effects on bud dormancy depth in Fagus sylvatica L.

The current state of knowledge on bud dormancy is limited. However, expanding such knowledge is crucial in order to properly model forest responses and feedback to future climate. Recent studies have shown that warming can decrease chilling accumulation and increase dormancy depth, thereby inducing delayed budburst in European beech (Fagus sylvatica L). Whether fall warming can advance spring phenology is unclear. To investigate the effect of warming on endodormancy of deciduous trees, we tested the impact of mild elevated temperature (+2.5-3.5 °C; temperature, on average, kept at 10 °C) in mid and late autumn on the bud dormancy depth and spring phenology of beech. We studied saplings by inducing periods of warming in greenhouses over a 2-year period. Even though warming reduced chilling accumulation in both years, we observed that the response of dormancy depth and spring budburst were year-specific. We found that warming during endodormancy peak could decrease the bud dormancy depth and therefore advance spring budburst. This effect appears to be modulated by factors such as the date of senescence onset and forcing intensity during endodormancy. Results from this study suggest that not only chilling but also forcing controls bud development during endodormancy and that extra forcing in autumn can offset reduced chilling.

Variability and limits of nitrogen and phosphorus resorption during foliar senescence.

Foliar nutrient resorption (NuR) plays a key role in ecosystem functioning and plant nutrient economy. Most of this recycling occurs during the senescence of leaves and is actively addressed by cells. Here, we discuss the importance of cell biochemistry, physiology, and subcellular anatomy to condition the outcome of NuR at the cellular level and to explain the existence of limits to NuR. Nutrients are transferred from the leaf in simple metabolites that can be loaded into the phloem. Proteolysis is the main mechanism for mobilization of N, whereas P mobilization requires the involvement of different catabolic pathways, making the dynamics of P in leaves more variable than those of N before, during, and after foliar senescence. The biochemistry and fate of organelles during senescence impose constraints that limit NuR. The efficiency of NuR decreases, especially in evergreen species, as soil fertility increases, which is attributed to the relative costs of nutrient acquisition from soil decreasing with increasing soil nutrient availability, while the energetic costs of NuR from senescing leaves remain constant. NuR is genetically determined, with substantial interspecific variability, and is environmentally regulated in space and time, with nutrient availability being a key driver of intraspecific variability in NuR.

Soil properties as key predictors of global grassland production: Have we overlooked micronutrients?

Fertilisation experiments have demonstrated that nutrient availability is a key determinant of biomass production and carbon sequestration in grasslands. However, the influence of nutrients in explaining spatial variation in grassland biomass production has rarely been assessed. Using a global dataset comprising 72 sites on six continents, we investigated which of 16 soil factors that shape nutrient availability associate most strongly with variation in grassland aboveground biomass. Climate and N deposition were also considered. Based on theory-driven structural equation modelling, we found that soil micronutrients (particularly Zn and Fe) were important predictors of biomass and, together with soil physicochemical properties and C:N, they explained more unique variation (32%) than climate and N deposition (24%). However, the association between micronutrients and biomass was absent in grasslands limited by NP. These results highlight soil properties as key predictors of global grassland biomass production and point to serial co-limitation by NP and micronutrients.

High-Resolution X-Ray Computed Tomography: A New Workflow for the Analysis of Xylogenesis and Intra-Seasonal Wood Biomass Production.

Understanding tree growth and carbon sequestration are of crucial interest to forecast the feedback of forests to climate change. To have a global understanding of the wood formation, it is necessary to develop new methodologies for xylogenesis measurements, valid across diverse wood structures and applicable to both angiosperms and gymnosperms. In this study, the authors present a new workflow to study xylogenesis using high-resolution X-ray computed tomography (HRXCT), which is generic and offers high potential for automatization. The HXRCT-based approach was benchmarked with the current classical approach (microtomy) on three tree species with contrasted wood anatomy (Pinus nigra, Fagus sylvatica, and Quercus robur). HRXCT proved to estimate the relevant xylogenesis parameters (timing, duration, and growth rates) across species with high accuracy. HRXCT showed to be an efficient avenue to investigate tree xylogenesis for a wide range of wood anatomies, structures, and species. HRXCT also showed its potential to provide quantification of intra-annual dynamics of biomass production through high-resolution 3D mapping of wood biomass within the forming growth ring.

Divergent responses of phenology and growth to summer and autumnal warming.

Plant phenology is highly sensitive to climate change, and shifts in autumnal foliar senescence are critical for plant productivity and nutrient cycling. Global warming has delayed the timing of foliar senescence, but the response of autumnal foliar senescence to nonuniform seasonal warming remains poorly understood, with experimental evidence in trees especially scarce. We therefore conducted a field experiment on seasonally asymmetric warming on 2-year-old larch (Larix principis-rupprechtii) seedlings in two hydrologically contrasting years (wet 2018 and dry 2019). Autumnal and year-round warming significantly delayed the timing of foliar senescence by 6 and 7 d in 2018, the wet year, with corresponding temperature sensitivities of 6.73 ± 1.47 and 8.26 ± 1.00 d/°C, respectively. Interestingly, the dates of senescence did not change across the warming treatments in 2019, the dry year. However, there was no significant effect of summer warming on the timing of foliar senescence neither in the wet nor dry year. The delayed autumnal foliar senescence was responsible for an increase in biomass only in the wet year, 2018. In contrast, summer warming, but not autumnal warming, increased the mortality of the seedlings in both 2018 and 2019. These results suggest that the hydrological conditions substantially modify the response of autumnal phenology and growth to seasonal warming. Autumnal warming increases growth, whereas summer warming could cause carbon starvation/hydraulic failure, reduce growth, and lead to higher mortality. Our results suggest that the functioning, ecosystem services, and sustainability of forests in the future depend on the strength and pattern of nonuniform seasonal warming. This study can inspire new research in phenology and tree growth in experiments with asymmetric warming.

Drought elicits contrasting responses on the autumn dynamics of wood formation in late successional deciduous tree species.

Research on wood phenology has mainly focused on reactivation of the cambium in spring. In this study we investigated if summer drought advances cessation of wood formation and if it has any influence on wood structure in late successional forest trees of the temperate zone. The end of xylogenesis was monitored between August and November in stands of European beech and pedunculate oak in Belgium for two consecutive years, 2017 and 2018, with the latter year having experienced an exceptional summer drought. Wood formation in oak was affected by the drought, with oak trees ceasing cambial activity and wood maturation about 3 weeks earlier in 2018 compared with 2017. Beech ceased wood formation before oak, but its wood phenology did not differ between years. Furthermore, between the 2 years, no significant difference was found in ring width, percentage of mature fibers in the late season, vessel size and density. In 2018, beech did show thinner fiber walls, whereas oak showed thicker walls. In this paper, we showed that summer drought can have an important impact on late season wood phenology xylem development. This will help to better understand forest ecosystems and improve forest models.

Timing of spring xylogenesis in temperate deciduous tree species relates to tree growth characteristics and previous autumn phenology.

We explored the timing of spring xylogenesis and its potential drivers in homogeneous mature forest stands in a temperate European region. Three species with contrasting leaf development dynamics and wood anatomy were studied: European beech, silver birch and pedunculate oak. Detailed phenological observations of xylogenesis and leaf phenology were performed from summer 2017 until spring 2018. Cambium reactivation (CR) occurred before the buds of oak and birch were swollen, whereas these two phenological phases were concurrent for beech. On the other hand, initial earlywood vessels were fully differentiated (FDIEV) after leaf unfolding for all three species. Timing of CR was correlated to average ring-width of the last 10 years (2008-17), tree diameter and, partially, with tree age. In addition, the timing of FDIEV was correlated to tree age and previous year's autumn phenology, i.e., timing of wood growth cessation and onset of leaf senescence. Multivariate models could explain up to 68% of the variability of CR and 55% of the variability of FDIEV. In addition to the 'species' factor, the variability could be explained by ca 30% by tree characteristics and previous year's autumn phenology for both CR and FDIEV. These findings are important to better identify which factors (other than environment) can be driving the onset of the growing season, and highlight the influence of tree growth characteristics and previous year's phenology on spring wood phenology, wood formation and, potentially, forest production.

Unaltered soil microbial community composition, but decreased metabolic activity in a semiarid grassland after two years of passive experimental warming.

Soil microbial communities regulate soil carbon feedbacks to climate warming through microbial respiration (i.e., metabolic rate). A thorough understanding of the responses of composition, biomass, and metabolic rate of soil microbial community to warming is crucial to predict soil carbon stocks in a future warmer climate. Therefore, we conducted a field manipulative experiment in a semiarid grassland on the Loess Plateau of China to evaluate the responses of the soil microbial community to increased temperature from April 2015 to December 2017. Soil temperature was 2.0°C higher relative to the ambient when open-top chambers (OTCs) were used. Warming did not affect microbial biomass or the composition of microbial functional groups. However, warming significantly decreased microbial respiration, directly resulting from soil pH decrease driven by the comediation of aboveground biomass increase, inorganic nitrogen increase, and moisture decrease. These findings highlight that the soil microbial community structure of semiarid grasslands resisted the short-term warming by 2°C, although its metabolic rate declined.

Inter-individual variability in spring phenology of temperate deciduous trees depends on species, tree size and previous year autumn phenology.

We explored the inter-individual variability in bud-burst and its potential drivers, in homogeneous mature stands of temperate deciduous trees. Phenological observations of leaves and wood formation were performed weekly from summer 2017 to summer 2018 for pedunculate oak, European beech and silver birch in Belgium. The variability of bud-burst was correlated to previous' year autumn phenology (i.e. the onset of leaf senescence and the cessation of wood formation) and tree size but with important differences among species. In fact, variability of bud-burst was primarily related to onset of leaf senescence, cessation of wood formation and tree height for oak, beech and birch, respectively. The inter-individual variability of onset of leaf senescence was not related to the tree characteristics considered and was much larger than the inter-individual variability in bud-burst. Multi-species multivariate models could explain up to 66% of the bud-burst variability. These findings represent an important advance in our fundamental understanding and modelling of phenology and tree functioning of deciduous tree species.

Modeling leaf senescence of deciduous tree species in Europe.

Autumnal leaf senescence signals the end of photosynthetic activities in temperate deciduous trees and consequently exerts a strong control on various ecological processes. Predicting leaf senescence dates (LSD) with high accuracy is thus a prerequisite for better understanding the climate-ecosystem interactions. However, modeling LSD at large spatial and temporal scales is challenging. In this study, first, we used 19972 site-year records (848 sites and four deciduous tree species) from the PAN European Phenology network to calibrate and evaluate six leaf senescence models during the period 1980-2013. Second, we extended the spatial analysis by repeating the procedure across Europe using satellite-derived end of growing season and a forest map. Overall, we found that models that considered photoperiod and temperature interactions outperformed models using simple temperature or photoperiod thresholds for Betula pendula, Fagus sylvatica and Quercus robur. On the contrary, no model displayed reasonable predictions for Aesculus hippocastanum. This inter-model comparison indicates that, contrary to expectation, photoperiod does not significantly modulate the accumulation of cooling degree days (CDD). On the other hand, considering the carryover effect of leaf unfolding date could promote the models' predictability. The CDD models generally matched the observed LSD at species level and its interannual variation, but were limited in explaining the inter-site variations, indicating that other environmental cues need to be considered in future model development. The discrepancies remaining between model simulations and observations highlight the need of manipulation studies to elucidate the mechanisms behind the leaf senescence process and to make current models more realistic.

Timeline of autumn phenology in temperate deciduous trees.

Cessation of xylem formation or wood growth (CWG) and onset of foliar senescence (OFS) are key autumn phenological events in temperate deciduous trees. Their timing is fundamental for the development and survival of trees, ecosystem nutrient cycling and the seasonal exchange of matter and energy between the biosphere and atmosphere, and affects the impact and feedback of forests to global change. A large-scale experimental effort and improved observational methods have allowed us to compare the timing of CWG and OFS for different deciduous tree species in Western Europe, particularly in silver birch, a pioneer species, and European beech, a late-succession species, at stands of different latitudes, of different levels of site fertility, for 2 years with contrasting meteorological and drought conditions, i.e., the low moderately dry 2017 and the extremely dry 2018. Specifically, we tested whether foliar senescence started before, after or concurrently with CWG. Onset of foliar senescence and CWG occurred generally between late September and early November, with larger differences across species and sites for OFS. Foliar senescence started concurrently with CWG in most cases, except for the drier 2018 and, for beech, at the coldest site, where OFS occurred significantly later than CWG. The behavior of beech in Spain, the southern edge of its European distribution, was unclear, with no CWG, but very low wood growth at the time of OFS. Our study suggests that OFS is generally triggered by the same drivers of CWG or when wood growth decreases in late summer, indicating an overarching mechanism of sink limitation as a possible regulator of the timing of foliar senescence.

Detecting the onset of autumn leaf senescence in deciduous forest trees of the temperate zone.

Information on the onset of leaf senescence in temperate deciduous trees and comparisons on its assessment methods are limited, hampering our understanding of autumn dynamics. We compare five field proxies, five remote sensing proxies and two data analysis approaches to assess leaf senescence onset at one main beech stand, two stands of oak and birch, and three ancillary stands of the same species in Belgium during 2017 and 2018. Across species and sites, onset of leaf senescence was not significantly different for the field proxies based on Chl leaf content and canopy coloration, except for an advanced canopy coloration during the extremely dry and warm 2018. Two remote sensing indices provided results fully consistent with the field data. A significant lag emerged between leaf senescence onset and leaf fall, and when a threshold of 50% change in the seasonal variable under study (e.g. Chl content) was used to derive the leaf senescence onset. Our results provide unprecedented information on the quality and applicability of different proxies to assess leaf senescence onset in temperate deciduous trees. In addition, a sound base is offered to select the most suited methods for the different disciplines that need this type of data.

Nutrient availability alters the correlation between spring leaf-out and autumn leaf senescence dates.

Leaf senescence (LS) affects tree fitness, species distribution and ecosystem structure and functioning. The drivers of LS and the processes underlying it have been studied, but the studies have mainly focused on environmental cues and have mainly been based on statistical analyses using in situ data sets. Experimental investigation and field verification of the processes and drivers are thus urgently needed. We conducted a nutrient-addition experiment after a spring-warming experiment in which an ~40-day range of leaf-out (LO) dates was induced in horse chestnut (Aesculus hippocastanum) and beech (Fagus sylvatica) saplings. We found that both increased nutrient supply and advanced LO date significantly affected the timing of LS, but their effects were opposite, as the former delayed and the latter advanced the senescence. The effects of nutrient supply and LO interacted species specifically. In chestnut, the delay of senescence caused by fertilization increased with the delay of LO and was thus stronger for individuals that flushed late in the spring. On the contrary, in beech the delay of senescence caused by fertilization decreased with the delay of LO and was insignificant for individuals with the latest LO. The experimental findings for beech were confirmed with mature trees at a regional scale. The interactive effect between nutrients and LO on senescence may be associated with variable sensitivity to photoperiod, growth sink limitation and/or direct effect of foliar nutrition on the timing of senescence. Our novel results show that the interactive effects of LO and nutrient supply on the timing of LS should be further addressed experimentally in forthcoming studies. It would also be interesting to consider our results in the further development of phenological models used in assessing the effects of climatic change. The differences found in the present study between horse chestnut and beech suggest that the results found for one species cannot necessarily be generalized to other species, so studies with different temperate tree species are called for.

Thinning Can Reduce Losses in Carbon Use Efficiency and Carbon Stocks in Managed Forests Under Warmer Climate.

Forest carbon use efficiency (CUE, the ratio of net to gross primary productivity) represents the fraction of photosynthesis that is not used for plant respiration. Although important, it is often neglected in climate change impact analyses. Here we assess the potential impact of thinning on projected carbon cycle dynamics and implications for forest CUE and its components (i.e., gross and net primary productivity and plant respiration), as well as on forest biomass production. Using a detailed process-based forest ecosystem model forced by climate outputs of five Earth System Models under four representative climate scenarios, we investigate the sensitivity of the projected future changes in the autotrophic carbon budget of three representative European forests. We focus on changes in CUE and carbon stocks as a result of warming, rising atmospheric CO2 concentration, and forest thinning. Results show that autotrophic carbon sequestration decreases with forest development, and the decrease is faster with warming and in unthinned forests. This suggests that the combined impacts of climate change and changing CO2 concentrations lead the forests to grow faster, mature earlier, and also die younger. In addition, we show that under future climate conditions, forest thinning could mitigate the decrease in CUE, increase carbon allocation into more recalcitrant woody pools, and reduce physiological-climate-induced mortality risks. Altogether, our results show that thinning can improve the efficacy of forest-based mitigation strategies and should be carefully considered within a portfolio of mitigation options.

Larger temperature response of autumn leaf senescence than spring leaf-out phenology.

Climate warming is substantially shifting the leaf phenological events of plants, and thereby impacting on their individual fitness and also on the structure and functioning of ecosystems. Previous studies have largely focused on the climate impact on spring phenology, and to date the processes underlying leaf senescence and their associated environmental drivers remain poorly understood. In this study, experiments with temperature gradients imposed during the summer and autumn were conducted on saplings of European beech to explore the temperature responses of leaf senescence. An additional warming experiment during winter enabled us to assess the differences in temperature responses of spring leaf-out and autumn leaf senescence. We found that warming significantly delayed the dates of leaf senescence both during summer and autumn warming, with similar temperature sensitivities (6-8 days delay per °C warming), suggesting that, in the absence of water and nutrient limitation, temperature may be a dominant factor controlling the leaf senescence in European beech. Interestingly, we found a significantly larger temperature response of autumn leaf senescence than of spring leaf-out. This suggests a possible larger contribution of delays in autumn senescence, than of the advancement in spring leaf-out, to extending the growing season under future warmer conditions.

Environmental impact assessment and monetary ecosystem service valuation of an ecosystem under different future environmental change and management scenarios; a case study of a Scots pine forest.

For a sustainable future, we must sustainably manage not only the human/industrial system but also ecosystems. To achieve the latter goal, we need to predict the responses of ecosystems and their provided services to management practices under changing environmental conditions via ecosystem models and use tools to compare the estimated provided services between the different scenarios. However, scientific articles have covered a limited amount of estimated ecosystem services and have used tools to aggregate services that contain a significant amount of subjective aspects and that represent the final result in a non-tangible unit such as 'points'. To resolve these matters, this study quantifies the environmental impact (on human health, natural systems and natural resources) in physical units and uses an ecosystem service valuation based on monetary values (including ecosystem disservices with associated negative monetary values). More specifically, the paper also focuses on the assessment of ecosystem services related to pollutant removal/generation flows, accounting for the inflow of eutrophying nitrogen (N) when assessing the effect of N leached to groundwater. Regarding water use/provisioning, evapotranspiration is alternatively considered a disservice because it implies a loss of (potential) groundwater. These approaches and improvements, relevant to all ecosystems, are demonstrated using a Scots pine stand from 2010 to 2089 for a combination of three environmental change and three management scenarios. The environmental change scenarios considered interannual climate variability trends and included alterations in temperature, precipitation, nitrogen deposition, wind speed, Particulate matter (PM) concentration and CO2 concentration. The addressed flows/ecosystem services, including disservices, are as follows: particulate matter removal, freshwater loss, CO2 sequestration, wood production, NOx emissions, NH3 uptake and nitrogen pollution/removal. The monetary ecosystem service valuation yields a total average estimate of 361-1242 euro ha(-1) yr(-1). PM2.5 (<2.5 µm) removal is the key service, with a projected value of 622-1172 euro ha(-1) yr(-1). Concerning environmental impact assessment, with net CO2 uptake being the most relevant contributing flow, a loss prevention of 0.014-0.029 healthy life years ha(-1) yr(-1) is calculated for the respective flows. Both assessment methods favor the use of the least intensive management scenario due to its resulting higher CO2 sequestration and PM removal, which are the most important services of the considered ones.

Modeled Changes in Potential Grassland Productivity and in Grass-Fed Ruminant Livestock Density in Europe over 1961-2010.

About 25% of European livestock intake is based on permanent and sown grasslands. To fulfill rising demand for animal products, an intensification of livestock production may lead to an increased consumption of crop and compound feeds. In order to preserve an economically and environmentally sustainable agriculture, a more forage based livestock alimentation may be an advantage. However, besides management, grassland productivity is highly vulnerable to climate (i.e., temperature, precipitation, CO2 concentration), and spatial information about European grassland productivity in response to climate change is scarce. The process-based vegetation model ORCHIDEE-GM, containing an explicit representation of grassland management (i.e., herbage mowing and grazing), is used here to estimate changes in potential productivity and potential grass-fed ruminant livestock density across European grasslands over the period 1961-2010. Here "potential grass-fed ruminant livestock density" denotes the maximum density of livestock that can be supported by grassland productivity in each 25 km × 25 km grid cell. In reality, livestock density could be higher than potential (e.g., if additional feed is supplied to animals) or lower (e.g., in response to economic factors, pedo-climatic and biotic conditions ignored by the model, or policy decisions that can for instance reduce livestock numbers). When compared to agricultural statistics (Eurostat and FAOstat), ORCHIDEE-GM gave a good reproduction of the regional gradients of annual grassland productivity and ruminant livestock density. The model however tends to systematically overestimate the absolute values of productivity in most regions, suggesting that most grid cells remain below their potential grassland productivity due to possible nutrient and biotic limitations on plant growth. When ORCHIDEE-GM was run for the period 1961-2010 with variable climate and rising CO2, an increase of potential annual production (over 3%) per decade was found: 97% of this increase was attributed to the rise in CO2, -3% to climate trends and 15% to trends in nitrogen fertilization and deposition. When compared with statistical data, ORCHIDEE-GM captures well the observed phase of climate-driven interannual variability in grassland production well, whereas the magnitude of the interannual variability in modeled productivity is larger than the statistical data. Regional grass-fed livestock numbers can be reproduced by ORCHIDEE-GM based on its simple assumptions and parameterization about productivity being the only limiting factor to define the sustainable number of animals per unit area. Causes for regional model-data misfits are discussed, including uncertainties in farming practices (e.g., nitrogen fertilizer application, and mowing and grazing intensity) and in ruminant diet composition, as well as uncertainties in the statistical data and in model parameter values.

Variation in leaf flushing date influences autumnal senescence and next year's flushing date in two temperate tree species.

Recent temperature increases have elicited strong phenological shifts in temperate tree species, with subsequent effects on photosynthesis. Here, we assess the impact of advanced leaf flushing in a winter warming experiment on the current year's senescence and next year's leaf flushing dates in two common tree species: Quercus robur L. and Fagus sylvatica L. Results suggest that earlier leaf flushing translated into earlier senescence, thereby partially offsetting the lengthening of the growing season. Moreover, saplings that were warmed in winter-spring 2009-2010 still exhibited earlier leaf flushing in 2011, even though the saplings had been exposed to similar ambient conditions for almost 1 y. Interestingly, for both species similar trends were found in mature trees using a long-term series of phenological records gathered from various locations in Europe. We hypothesize that this long-term legacy effect is related to an advancement of the endormancy phase (chilling phase) in response to the earlier autumnal senescence. Given the importance of phenology in plant and ecosystem functioning, and the prediction of more frequent extremely warm winters, our observations and postulated underlying mechanisms should be tested in other species.

The impact of winter and spring temperatures on temperate tree budburst dates: results from an experimental climate manipulation.

Budburst phenology is a key driver of ecosystem structure and functioning, and it is sensitive to global change. Both cold winter temperatures (chilling) and spring warming (forcing) are important for budburst. Future climate warming is expected to have a contrasting effect on chilling and forcing, and subsequently to have a non-linear effect on budburst timing. To clarify the different effects of warming during chilling and forcing phases of budburst phenology in deciduous trees, (i) we conducted a temperature manipulation experiment, with separate winter and spring warming treatments on well irrigated and fertilized saplings of beech, birch and oak, and (ii) we analyzed the observations with five temperature-based budburst models (Thermal Time model, Parallel model, Sequential model, Alternating model, and Unified model). The results show that both winter warming and spring warming significantly advanced budburst date, with the combination of winter plus spring warming accelerating budburst most. As expected, all three species were more sensitive to spring warming than to winter warming. Although the different chilling requirement, the warming sensitivity was not significantly different among the studied species. Model evaluation showed that both one- and two- phase models (without and with chilling, respectively) are able to accurately predict budburst. For beech, the Sequential model reproduced budburst dates best. For oak and birch, both Sequential model and the Thermal Time model yielded good fit with the data but the latter was slightly better in case of high parameter uncertainty. However, for late-flushing species, the Sequential model is likely be the most appropriate to predict budburst data in a future warmer climate.