Effectiveness of freezing temperatures on dormancy release of temperate woody species.

BACKGROUND AND AIMS: Spring phenological change of plants in response to global warming may affect many ecological processes and functions. Chilling temperature regulates budburst date by releasing dormancy. However, whether freezing temperatures (<0 °C) contribute to dormancy release remains of debate. Our poor understanding of the role of chilling makes estimating shifts in budburst date difficult. METHODS: A 2-year chilling-forcing experiment was explicitly designed to test the effects of chilling temperatures on dormancy release of nine temperate woody species in Beijing, China. A total of 1620 twigs were first exposed to a wide range of temperatures (-10 to 10 °C) with different durations and then moved to growth chambers. Based on budburst data in experimental conditions, we examined whether freezing temperatures are effective on dormancy release. We also developed a new framework for constructing chilling functions based on the curve between chilling duration and forcing requirement (FR) of budburst. The chilling function derived from this framework was not affected by experimental forcing conditions. KEY RESULTS: We demonstrated that freezing temperatures down to -10 °C were effective in dormancy release. The rate of dormancy release, indicated by the rate of decay in the chilling duration-FR curve, did not differ significantly between chilling temperatures in most cases, although it exhibited a maximum value at 0 or 5 °C. The chilling function-associated phenological models could simulate budburst date from independent experimental and observational data with a mean RMSE of 7.07 d. CONCLUSIONS: The effective freezing temperatures found here are contrary to the well-known assumption of <0 °C temperature generally not contributing to accumulated chilling in many previous chilling functions. A chilling function assuming that temperature below an upper temperature threshold has the same effects on dormancy release could be adopted to calculate chilling accumulation when using experiments to develop spring phenological models based on the chilling-forcing relationship.

Widespread decline in winds delayed autumn foliar senescence over high latitudes.

The high northern latitudes (>50°) experienced a pronounced surface stilling (i.e., decline in winds) with climate change. As a drying factor, the influences of changes in winds on the date of autumn foliar senescence (DFS) remain largely unknown and are potentially important as a mechanism explaining the interannual variability of autumn phenology. Using 183,448 phenological observations at 2,405 sites, long-term site-scale water vapor and carbon dioxide flux measurements, and 34 y of satellite greenness data, here we show that the decline in winds is significantly associated with extended DFS and could have a relative importance comparable with temperature and precipitation effects in contributing to the DFS trends. We further demonstrate that decline in winds reduces evapotranspiration, which results in less soil water losses and consequently more favorable growth conditions in late autumn. In addition, declining winds also lead to less leaf abscission damage which could delay leaf senescence and to a decreased cooling effect and therefore less frost damage. Our results are potentially useful for carbon flux modeling because an improved algorithm based on these findings projected overall widespread earlier DFS than currently expected by the end of this century, contributing potentially to a positive feedback to climate.

Winter warming offsets one half of the spring warming effects on leaf unfolding.

Winter temperature-related chilling and spring temperature-related forcing are two major environmental cues shaping the leaf-out date of temperate species. To what degree insufficient chilling caused by winter warming would slow phenological responses to spring warming remains unclear. Using 27,071 time series of leaf-out dates for 16 tree species in Europe, we constructed a phenological model based on the linear or exponential function between the chilling accumulation (CA) and forcing requirements (FR) of leaf-out. We further used the phenological model to quantify the relative contributions of chilling and forcing on past and future spring phenological change. The results showed that the delaying effect of decreased chilling on the leaf-out date was prevalent in natural conditions, as more than 99% of time series exhibited a negative relationship between CA and FR. The reduction in chilling linked to winter warming from 1951 to 2014 could offset about one half of the spring phenological advance caused by the increase in forcing. In future warming scenarios, if the same model is used and a linear, stable correlation between CA and FR is assumed, declining chilling will continuously offset the advance of leaf-out to a similar degree. Our study stresses the importance of assessing the antagonistic effects of winter and spring warming on leaf-out phenology.

Varying temperature sensitivity of bud-burst date at different temperature conditions.

The relationship between the rate of development (DR) of bud-burst and temperature may be nonlinear, which could lead to varying temperature sensitivity (TS) of budburst date under different climate conditions. In order to determine the functional form between DR/TS and temperature, we gathered twigs with flower buds of five woody plants (Malus halliana, Forsythia suspense, Crataegus pinnatifida, Prunus cerasifera F. atropurpurea, and Berberis thunbergii var. atropurpurea Chenault) in early spring of 2017 at Beijing, and placed them in six growth chambers at same daylength but different temperature conditions (5, 10, 15, 20, 25, and 30 °C). The proportion of bud- burst was recorded every 2 or 3 days for each species at each temperature condition. The results showed that the proportion of bud-burst followed the logistic function over time at a given temperature. Subsequently, we developed a mathematical model to simulate the proportion of bud-burst at any temperature and date. The DR and TS were parameterized using a differential method. The simulation results showed that the DR increased monotonically with the rise in temperature, but only two species could reach the maximum value at 30 °C. The TS decreased with the increase in temperature, but this effect was weak when the temperature was high enough. These findings suggested that the predicted warming in the future may result in a slowdown in the advance of spring phenology of woody plants.

Comparison of chilling and heat requirements for leaf unfolding in deciduous woody species in temperate and subtropical China.

Climate warming has advanced the spring phenology of many plant species by accelerating heat accumulation. However, delayed phenophases due to insufficient chilling have also been reported. Based on phenological observation data (1963-2010), we compared the effects of preseason chill and heat accumulation on leaf unfolding dates of four deciduous woody species (Lagerstroemia indica, Robinia pseudoacacia, Sophora japonica, and Ulmus pumila) in temperate and subtropical regions of China. Daily chill and heat accumulation were calculated by two chilling models (the Positive Utah Model and the Dynamic Model) and the Growing Degree Hour (GDH) Model. We determined the temporal trends in chill and heat accumulations for leaf unfolding of the four species. The results showed that there were shorter chilling periods in the subtropics than in temperate sites because the chilling period typically started later and ended earlier. There was no significant difference in the length of the forcing period in the different regions. The chilling requirements for leaf unfolding were higher in temperate regions (1344.9-1798.9 chilling units (CU) or 64.7-79.4 chilling portions (CP)) than in the subtropics (1145.9-1828.1 CU or 47.9-75.2 CP). Plants in the subtropics needed higher forcing temperatures (4135.8-10084.8 GDH) than those in temperate regions (3292.0-8383.6 GDH). The earlier-leafing species (e.g., U. pumila) had a lower heat requirement for leaf unfolding than the later-leafing species (e.g., L. indica). A significant increase in heat accumulation was found at all sites except Guiyang, while chill accumulation only increased in Beijing.

Trends in maize (Zea mays L.) phenology and sensitivity to climate factors in China from 1981 to 2010.

Changes in crop phenology may reflect crop responses and adaptation to climate change. In this study, we used observational data (1981-2010) of maize (Zea mays L.) phenology from agricultural meteorological stations in the major maize-growing regions of China to examine spatiotemporal changes in the phenologies and growth periods and associated sensitivities to changes in major climatic factors. The results showed that, during the study period, sowing, tasseling, and maturity dates for maize were delayed in most maize growth regions. The lengths of vegetative growth period (VGP, from emergence to tasseling) were increased in spring and spring-summer maize growth regions and decreased in summer maize growth regions; the lengths of the maize reproductive growth period (RGP, from tasseling to maturity) and whole growth period (WGP, from emergence to maturity) were mostly extended (except NWMR_SU). Overall, sensitivity of maize VGP, RGP, and WGP was negatively related to average temperature (P < 0.01) and positively related to precipitation and sunshine hours (P < 0.01); there were variations in sensitivity among regions and data station locations. Precipitation was a driver of growth period length in the northwest inland maize region, whereas mean temperature and sunshine hours were drivers in the southwest hilly region.

Changes in flowering phenology of woody plants from 1963 to 2014 in North China.

Existing evidence demonstrates that the first flowering date (FFD) of most plant species became earlier in response to temperature increase over the past several decades. However, the studies on changes in flowering duration (FD) were limited. By using the non-parametric Theil-Sen estimator, this study investigated the temporal trends in 127 time series of FFD, end of flowering date (EFD), and FD of 97 woody plants from 1963 to 2014 at three sites (Harbin, Beijing, and Xi'an) in North China. The relationship between flowering phenophases and temperature was analyzed using two phenological models. The results showed that most of FFD and EFD time series exhibited an apparent advancing trend. Among them, trends of 52.0% (40.9%) of FFD (EFD) time series were significant (P < 0.05). FFD and EFD time series (95.3 and 89.8%, respectively) responded negatively and significantly to preseason temperature (P < 0.05). The direction of FD changes varied among sites and species. On average, a shortening trend of FD was observed at Harbin (-0.51 days decade-1), with 7.5% of species significantly. However, FD on average extended by 0.42 and 0.93 days decade-1 at Beijing (24.5% significantly) and Xi'an (28.9% significantly), respectively. The regression models could simulate the interannual changes in FFD and EFD with the mean goodness of fit (R2) ranging from 0.37 to 0.67, but fail to simulate the changes in FD accurately (R2 ranging from 0.09 to 0.18). The growing degree day model could improve the R2 for simulating FFD and EFD except for FD. Therefore, more phenological models need to be tested, and more drivers of FD need to be further investigated.

Variations in the temperature sensitivity of spring leaf phenology from 1978 to 2014 in Mudanjiang, China.

Continuous long-term temperature sensitivity (ST) of leaf unfolding date (LUD) and main impacting factors in spring in the period 1978-2014 for 40 plant species in Mudanjiang, Heilongjiang Province, Northeast China, were analyzed by using observation data from the China Phenological Observation Network (CPON), together with the corresponding meteorological data from the China Meteorological Data Service Center. Temperature sensitivities, slopes of the regression between LUD and mean temperature during the optimum preseason (OP), were analyzed using 15-year moving window to determine their temporal trends. Major factors impacting ST were then chosen and evaluated by applying a random sampling method. The results showed that LUD was sensitive to mean temperature in a defined period before phenophase onset for all plant species analyzed. Over the period 1978-2014, the mean ST of LUD for all plant species was - 3.2 ± 0.49 days °C-1. The moving window analysis revealed that 75% of species displayed increasing ST of LUD, with 55% showing significant increases (P < 0.05). ST for the other 25% exhibited a decreasing trend, with 17% showing significant decreases (P < 0.05). On average, ST increased by 16%, from - 2.8 ± 0.83 days °C-1 during 1980-1994 to - 3.30 ± 0.65 days °C-1 during 2000-2014. For species with later LUD and longer OP, ST tended to increase more, while species with earlier LUD and shorter OP tended to display a decreasing ST. The standard deviation of preseason temperature impacted the temporal variation in ST. Chilling conditions influenced ST for some species, but photoperiod limitation did not have significant or coherent effects on changes in ST.

Impacts of global warming on phenology of spring leaf unfolding remain stable in the long run.

The impact of spring temperature forcing on the timing of leaf unfolding of plants (temperature sensitivity, ST) is one important indicator of how and to what degree plant species track climate change. Fu et al. (Nature 526:104-107, 2015) found that ST has significantly decreased from the 1980-1994 to the 1999-2013 period for seven mid-latitude tree species in Europe. However, long-term changes in ST over the past 60 years are still not clear. Here, using in situ observations of leaf unfolding for seven dominant European tree species, we analyze the temporal change in ST over decadal time scales extending the data series back to 1951. Our results demonstrate that ST shows no statistically significant change within shifting 30-year windows from 1951 to 2013 and remains stable between 1951-1980 and 1984-2013 (3.6 versus 3.7 days °C-1). This result suggests that the significant decrease in ST over the past 33 years could not be sustained when examining the trends of phenological responses in the long run. Therefore, we could not conclude that tree spring phenology advances will slow down in the future, and the ST changes in warming scenarios are still uncertain.

Reply to communications by Fu et al. international journal of biometeorology.

Temperature sensitivity of plant phenology (ST) is a determining factor of as to what degree climate change impacts on plant species. Fu et al . (Int J Biometeorol 60:1611-1613, 2016) claimed that long long-term linear trends mask phenological shifts. However, the decreased and increased ST was both found in warming scenarios. The conceptual scheme telling the nonlinear relationship between spring temperature and leaf unfolding date proposed by Fu et al . (Int J Biometeorol 60:1611-1613, 2016) cannot be supported by observation data across Europe. Therefore, linking declined ST to climate warming is misleading, and future ST changes are more uncertain than they suggested.

Phenological response to climate change in China: a meta-analysis.

The change in the phenology of plants or animals reflects the response of living systems to climate change. Numerous studies have reported a consistent earlier spring phenophases in many parts of middle and high latitudes reflecting increasing temperatures with the exception of China. A systematic analysis of Chinese phenological response could complement the assessment of climate change impact for the whole Northern Hemisphere. Here, we analyze 1263 phenological time series (1960-2011, with 20+ years data) of 112 species extracted from 48 studies across 145 sites in China. Taxonomic groups include trees, shrubs, herbs, birds, amphibians and insects. Results demonstrate that 90.8% of the spring/summer phenophases time series show earlier trends and 69.0% of the autumn phenophases records show later trends. For spring/summer phenophases, the mean advance across all the taxonomic groups was 2.75 days decade(-1) ranging between 2.11 and 6.11 days decade(-1) for insects and amphibians, respectively. Herbs and amphibians show significantly stronger advancement than trees, shrubs and insect. The response of phenophases of different taxonomic groups in autumn is more complex: trees, shrubs, herbs and insects show a delay between 1.93 and 4.84 days decade(-1), while other groups reveal an advancement ranging from 1.10 to 2.11 days decade(-1) . For woody plants (including trees and shrubs), the stronger shifts toward earlier spring/summer were detected from the data series starting from more recent decades (1980s-2000s). The geographic factors (latitude, longitude and altitude) could only explain 9% and 3% of the overall variance in spring/summer and autumn phenological trends, respectively. The rate of change in spring/summer phenophase of woody plants (1960s-2000s) generally matches measured local warming across 49 sites in China (R=-0.33, P<0.05).

Geographical pattern in first bloom variability and its relation to temperature sensitivity in the USA and China.

Advance in spring plant phenology over the last several decades has been found in all continents of the Northern Hemisphere. Compared to the studies detecting phenological trends, the studies investigating the geographical pattern of phenological variability (including mean date and magnitude of variability) are rather limited. In this study, we analyzed spatial pattern of mean date and standard deviation (SD) of first bloom date (FBD) time series (≥15 years) for black locust (Robinia pseudoacacia) at 22 stations in China, common lilac (Syringa vulgaris) at 79 stations in the Western US and Chinese lilac (Syringa chinensis) at 45 stations in the Eastern US. Subsequently, the impact of geographical factors (latitude, longitude, and altitude) on the mean date and SD was quantified by using the multiple regression analysis method. Meanwhile, the relationship between FBD variability and temperature sensitivity of FBD was examined. Results showed that the mean FBD highly depended on geographical factors for all the three species. Compared to the mean date, the dependence of SD of FBD time series on geographical factors was weaker. The geographical factors could only explain 13 to 31 % of spatial variance in SD of FBD. The negative regression coefficients of latitude (P < 0.05 except black locust) indicated that FBD is more variable at lower latitude. At most of stations, significant and negative correlations between FBD and preseason temperature on interannual scale were found, but the temperature sensitivity varied among different stations. The magnitude of temperature sensitivity decreased with increasing latitude. In general, the locations at lower latitude had earlier and more variable spring phenophase and showed stronger phenological response to climate change than the locations at higher latitude.

The spatial pattern of leaf phenology and its response to climate change in China.

Leaf phenology has been shown to be one of the most important indicators of the effects of climate change on biological systems. Few such studies have, however, been published detailing the relationship between phenology and climate change in Asian contexts. With the aim of quantifying species' phenological responsiveness to temperature and deepening understandings of spatial patterns of phenological and climate change in China, this study analyzes the first leaf date (FLD) and the leaf coloring date (LCD) from datasets of four woody plant species, Robinia pseudoacacia, Ulmus pumila, Salix babylonica, and Melia azedarach, collected from 1963 to 2009 at 47 Chinese Phenological Observation Network (CPON) stations spread across China (from 21° to 50° N). The results of this study show that changes in temperatures in the range of 39-43 days preceding the date of FLD of these plants affected annual variations in FLD, while annual variations in temperature in the range of 71-85 days preceding LCD of these plants affected the date of LCD. Average temperature sensitivity of FLD and LCD for these plants was -3.93 to 3.30 days °C(-1) and 2.11 to 4.43 days °C⁻¹, respectively. Temperature sensitivity of FLD was found to be stronger at lower latitudes or altitude as well as in more continental climates, while the response of LCD showed no consistent pattern. Within the context of significant warming across China during the study period, FLD was found to have advanced by 5.44 days from 1960 to 2009; over the same period, LCD was found to have been delayed by 4.56 days. These findings indicate that the length of the growing season of the four plant species studied was extended by a total of 10.00 days from 1960 to 2009. They also indicate that phenological response to climate is highly heterogeneous spatially.

Simulating changes in the leaf unfolding time of 20 plant species in China over the twenty-first century.

Recent shifts in phenology reflect the biological response to current climate change. Aiming to enhance our understanding of phenological responses to climate change, we developed, calibrated and validated spatio-temporal models of first leaf date (FLD) for 20 broadleaved deciduous plants in China. Using daily meteorological data from the Chinese Meteorological Administration and the Community Climate System Model, version 3 (CCSM3) created using three IPCC scenarios (A2, A1B and B1), we described the FLD time series of each species over the past 50 years, extrapolating from these results to simulate estimated FLD changes for each species during the twenty-first century. Model validation suggests that our spatio-temporal models can simulate FLD accurately with R² (explained variance) >0.60. Model simulations show that, from 1952 to 2007, the FLD in China advanced at a rate of -1.14 days decade⁻¹) on average. Furthermore, changes in FLD showed noticeable variation between regions, with clearer advances observed in the north than in the south of the country. The model indicates that the advances in FLD observed from 1952-2007 in China will continue over the twenty-first century, although significant differences among species and different climate scenarios are expected. The average trend of FLD advance in China during the twenty-first century is modeled as being -1.92 days decade⁻¹ under the A2 scenario, -1.10 days decade⁻¹ under the A1B scenario and -0.74 days decade⁻¹ under the B2 scenario. The spatial pattern of FLD change for the period 2011-2099 is modeled as being similar but showing some difference from patterns in the 1952-2007 period. At the interspecific level, early-leafing species were found to show a greater advance in FLD, while species with larger distributions tended to show a weaker advance in FLD. These simulated changes in phenology may have significant implications for plant distribution as well as ecosystem structure and function.

Spring wood phenology responds more strongly to chilling temperatures than bud phenology in European conifers.

A comparative assessment of bud and wood phenology could aid a better understanding of tree growth dynamics. However, the reason for asynchronism or synchronism in leaf and cambial phenology remains unclear. To test the assumption that the temporal relationship between the budburst date and the onset date of wood formation is due to their common or different responses to environmental factors, we constructed a wood phenology dataset from previous literature, and compared it with an existing bud phenology dataset in Europe. We selected three common conifers (Larix decidua Mill., Picea abies (L.) H. Karst. and Pinus sylvestris L.) in both datasets and analyzed 909 records of the onset of wood formation at 47 sites and 238,720 records of budburst date at 3051 sites. We quantified chilling accumulation (CA) and forcing requirement (FR) of budburst and onset of wood formation based on common measures of CA and FR. We then constructed negative exponential CA-FR curves for bud and wood phenology separately. The results showed that the median, variance and probability distribution of CA-FR curves varied significantly between bud and wood phenology for three conifers. The different FR under the same chilling condition caused asynchronous bud and wood phenology. Furthermore, the CA-FR curves manifested that wood phenology was more sensitive to chilling than bud phenology. Thus, the FR of the onset of wood formation increases more than that of budburst under the same warming scenarios, explaining the stronger earlier trends in the budburst date than the onset date of woody formation simulated by the process-based model. Our work not only provides a possible explanation for asynchronous bud and wood phenology from the perspective of organ-specific responses to chilling and forcing, but also develops a phenological model for predicting both bud and wood phenology with acceptable uncertainties.

Multiple phenological responses to climate change among 42 plant species in Xi'an, China.

Phenological data of 42 woody plants in a temperate deciduous forest from the Chinese Phenological Observation Network (CPON) and the corresponding meteorological data from 1963 to 2011 in Xi'an, Shaanxi Province, China were collected and analyzed. The first leaf date (FLD), leaf coloring date (LCD) and first flower date (FFD) are revealed as strong biological signals of climatic change. The FLD, LCD and FFD of most species are sensitive to average temperature during a certain period before phenophase onset. Regional precipitation also has a significant impact on phenophases of about half of the species investigated. Affected by climate change, the FLD and FFD of these species have advanced by 5.54 days and 10.20 days on average during 2003-2011 compared with the period 1963-1996, respectively. Meanwhile, the LCD has delayed by 10.59 days, and growing season length has extended 16.13 days. Diverse responses of phenology commonly exist among different species and functional groups during the study period. Especially for FFD, the deviations between the above two periods ranged from -20.68 to -2.79 days; biotic pollination species showed a significantly greater advance than abiotic pollination species. These results were conducive to the understanding of possible changes in both the structure of plant communities and interspecific relationships in the context of climate change.

Magnitude and direction of green-up date in response to drought depend on background climate over Mongolian grassland.

Increasing drought is one major consequence of ongoing global climate change and is expected to cause significant changes in vegetation phenology, especially for naturally vulnerable ecosystems such as grassland. However, the linkage between the response characteristic of green-up date (GUD) to drought and background climate remains largely unknown. Here, we focused on how the GUD of Mongolian grassland responds to extreme drought events (EDE). We first extracted the GUD from the MODIS Enhanced Vegetation Index data during 2001-2020 and identified the preseason EDE using the standardized precipitation evapotranspiration index data. Subsequently, we quantified the response of GUD to preseason EDE (DGUD) in each pixel as the difference in GUD between drought and normal years. The effect of 12 factors on DGUD was analyzed using the random forest algorithm. The results showed that the GUD under EDE may delay or advance by > 20 days compared to normal years. For the regions with mean annual temperature > -2 °C, the GUD was delayed under EDE due to the dominant role of water restriction on GUD, while the GUD was advanced under EDE in colder areas due to the warmer temperature during drought. However, the magnitude of delay in GUD under drought was greater in regions with less precipitation and more severe droughts. Our results could help to develop appropriate management strategies to mitigate the impacts of drought on grasslands.

Autumn phenology of tree species in China is associated more with climate than with spring phenology and phylogeny.

Both biotic and abiotic factors restrict changes in autumn phenology, yet their effects remain ambiguous, which hinders the accurate prediction of phenology under future climate change. In this study, based on the phenological records of 135 tree species at ten sites in China during 1979-2018, we first investigated the effects of climatic factors (temperature, precipitation, insolation and wind speed) and spring phenology on interannual changes in leaf coloring date (LCD) with the partial correlation analysis, and assessed the relative importance of phylogeny and native climate to LCD differences among species by using multivariate regression and phylogenetic eigenvector regression approach. The results showed that the effects of climate factors on interannual changes in LCD were more significant than spring phenology. In general, temperature played a more important role in cold regions (e.g. the northeast region), while the control of insolation on LCD was stronger in the warmer and wetter regions (e.g. the north, east and southwest regions). In addition, the effects of precipitation and wind speed were more evident in arid regions (e.g. the northwest region). We also found considerable effects of both native climate and phylogeny on the LCD differences among species, despite the contribution of native climate being almost 2~5 times greater than that of the phylogeny. Our findings confirmed and quantified the combined effects of climate, spring phenology and phylogeny on the autumn phenology of plants, which could help better understand the driving factors and influencing mechanism of plant phenology and provide a reference for the calibration and optimization of phenological models.

Modeling the effect of adaptation to future climate change on spring phenological trend of European beech (Fagus sylvatica L.).

Temperate trees could cope with climate change through phenotypic plasticity of phenological key events or adaptation in situ via selection on genetic variation. However, the relative contribution of local adaptation and phenotypic plasticity to phenological change is unclear for many ecologically important tree species. Here, we analyzed the leaf-out data of European beech (Fagus sylvatica L.) from 50 provenances planted in 7 trial sites. We first constructed a function between chilling accumulation (CA) and photoperiod-associated heat requirement (PHR) of leaf-out date for each provenance and quantified the relationship between parameters of the CA-PHR function and climatic variables at provenance origins by using the random forest model. Furthermore, we used the provenance-specific CA-PHR function to simulate future leaf-out dates under two climate change scenarios (RCP 4.5 and 8.5) and two assumptions (no adaptation and adaptation). The results showed that both CA, provenance, and their interactions affected the PHR of leaf-out. The provenances from southeastern Europe exhibited a stronger response of PHR to CA and thus flushed earlier than northwestern provenances. The parameters of the CA-PHR function were connected with climatic variables (e.g., mean diurnal temperature range, temperature seasonality) at the originating sites of each provenance. If only considering the phenotypic plasticity, the leaf-out date of European beech in 2070-2099 will advance by 6.8 and 9.0 days on average relative to 1951-2020 under RCP 4.5 and RCP 8.5, respectively. However, if F. sylvatica adapts to future climate change by adopting the current strategy, the advance of the leaf-out date will weaken by 1.4 and 3.4 days under RCP 4.5 and RCP 8.5, respectively. Our results suggest that the European beech could slow down its spring phenological advances and reduce its spring frost risk if it adopts the current strategy to adapt to future climate change.

Low temperature and short daylength interact to affect the leaf senescence of two temperate tree species.

Temperature and photoperiod are two major environmental cues shaping the leaf senescence of temperate tree species. However, how the control of leaf senescence is split between photoperiod and temperature is unknown for many ecologically important species. Here, we conducted a growth chamber experiment to test the effects of temperature (6, 9, 18 and 21°C) and photoperiod (8 and 16 h daylength) on leaf senescence of two temperate tree species (Quercus mongolica Fisch. and Larix principis-rupprechtii Mayr.) distributed in montane forest of China. The results showed that low temperature (LT) alone could induce leaf senescence of both species under long daylength (LD) conditions, but the leaf senescence of L. principis-rupprechtii was more sensitive to the decrease in temperature than that of Q. mongolica under the LD condition. Short daylength (SD) alone could only induce the leaf senescence of L. principis-rupprechtii, suggesting that the photoperiod sensitivity varies between species. SD could accelerate the LT-induced senescence, but the effect of SD reduced with the decrease in temperature. Based on these findings, we developed a new autumn phenology model by incorporating interspecific differences in the photoperiod sensitivity of leaf senescence. Compared with the three existing process-based autumn phenology models, the new model was more robust in simulating the experimental data. When employing these models to available long-term phenological data, our new model also performed best in reproducing the observed leaf senescence date of two closely related species (Quercus robur L. and Larix decidua Mill.). These results enhance our understanding of how LT and SD control leaf senescence. The prediction of the climate change impacts on forest carbon uptake could be improved by incorporating this new autumn phenological model into the terrestrial biosphere models.