Field evidence for earlier leaf-out dates in alpine grassland on the eastern Tibetan Plateau from 1990 to 2006

Worldwide, many plant species are experiencing an earlier onset of spring phenophases due to climate warming. Rapid recent temperature increases on the Tibetan Plateau (TP) have triggered changes in the spring phenology of the local vegetation. However, remote sensing studies of the land surface phenology have reached conflicting interpretations about green-up patterns observed on the TP since the mid-1990s. We investigated this issue using field phenological observations from 1990 to 2006, for 11 dominant plants on the TP at the levels of species, families (Gramineae—grasses and Cyperaceae—sedges) and vegetation communities (alpine meadow and alpine steppe). We found a significant trend of earlier leaf-out dates for one species (Koeleria cristata). The leaf-out dates of both Gramineae and Cyperaceae had advanced (the latter significantly, starting an average of 9 days later per year than the former), but the correlation between them was significant. The leaf-out dates of both vegetation communities also advanced, but the pattern was only significant in the alpine meadow. This study provides the first field evidence of advancement in spring leaf phenology on the TP and suggests that the phenology of the alpine steppe can differ from that of the alpine meadow. These findings will be useful for understanding ecosystem responses to climate change and for grassland management on the TP.     

Predicted responses of arctic and alpine ecosystems to altered seasonality under climate change

Global climate change is already having significant impacts on arctic and alpine ecosystems, and ongoing increases in temperature and altered precipitation patterns will affect the strong seasonal patterns that characterize these temperature‐limited systems. The length of the potential growing season in these tundra environments is increasing due to warmer temperatures and earlier spring snow melt. Here, we compare current and projected climate and ecological data from 20 Northern Hemisphere sites to identify how seasonal changes in the physical environment due to climate change will alter the seasonality of arctic and alpine ecosystems. We find that although arctic and alpine ecosystems appear similar under historical climate conditions, climate change will lead to divergent responses, particularly in the spring and fall shoulder seasons. As seasonality changes in the Arctic, plants will advance the timing of spring phenological events, which could increase plant nutrient uptake, production, and ecosystem carbon (C) gain. In alpine regions, photoperiod will constrain spring plant phenology, limiting the extent to which the growing season can lengthen, especially if decreased water availability from earlier snow melt and warmer summer temperatures lead to earlier senescence. The result could be a shorter growing season with decreased production and increased nutrient loss. These contrasting alpine and arctic ecosystem responses will have cascading effects on ecosystems, affecting community structure, biotic interactions, and biogeochemistry.     

高寒草甸植物群落生长发育特征与气候因子的关系

为合理利用高寒草甸资源，探讨近年来气候变化对高寒草甸的影响，以青海省甘德县高寒草甸为例，基于牧业气象站1976-2006年的气象资料和1994-2006年的牧草观测资料，分析了草地植被地上生物量、高度、盖度和物候期等群落特征以及当地气温、降雨等气象因素的年际变化趋势，采用典型相关分析法和逐步回归分析法对草地植物群落特征变化与气象因子的关系进行了研究，综合分析了影响植被生长状况的关键因子，结果表明：（1）青藏高原高寒草甸总体呈年均气温和平均地温上升、年降水量下降的"暖干化"趋势，牧草盖度高度增大，产量减少，整体观测水平下的牧草物候期推迟。（2）牧草的高度、盖度及产量对不同气候因子的响应程度不同。牧草高度与盖度对温度因子的变化更敏感，牧草产量对水分因子的变化更敏感。平均地温和相对湿度越高，牧草高度越高，产量越多。（3）不同牧草的物候期受不同气象因子的影响，变化趋势也不相同。从整体水平上看，牧草物候期对温度因子更敏感，温度越高，物候期越提前。     

The influence of local spring temperature variance on temperature sensitivity of spring phenology

The impact of climate warming on the advancement of plant spring phenology has been heavily investigated over the last decade and there exists great variability among plants in their phenological sensitivity to temperature. However, few studies have explicitly linked phenological sensitivity to local climate variance. Here, we set out to test the hypothesis that the strength of phenological sensitivity declines with increased local spring temperature variance, by synthesizing results across ground observations. We assemble ground‐based long‐term (20–50 years) spring phenology database (PEP725 database) and the corresponding climate dataset. We find a prevalent decline in the strength of phenological sensitivity with increasing local spring temperature variance at the species level from ground observations. It suggests that plants might be less likely to track climatic warming at locations with larger local spring temperature variance. This might be related to the possibility that the frost risk could be higher in a larger local spring temperature variance and plants adapt to avoid this risk by relying more on other cues (e.g., high chill requirements, photoperiod) for spring phenology, thus suppressing phenological responses to spring warming. This study illuminates that local spring temperature variance is an understudied source in the study of phenological sensitivity and highlight the necessity of incorporating this factor to improve the predictability of plant responses to anthropogenic climate change in future studies.     

Effects of experimental warming on plant reproductive phenology in Xizang alpine meadow北大核心CSCD

Aims: Climate warming strongly influences reproductive phenology of plants in alpine and arctic ecosystems. Here we focus on phenological shifts caused by warming in a typical alpine meadow on the Qinghai-Xizang Plateau. Our objective was to explore phenological responses of alpine plant species to experimental warming. Methods: Passive warming was achieved using open-top chambers (OTCs). The treatments included control (C), and four levels of warming (T1, T2, T3, T4). We selected Kobresia pygmaea, Potentilla saundersiana, Potentilla cuneata, Stipa purpurea, Festuca coelestis and Youngia simulatrix as the focal species. Plant phenology was scored every 3-5 days in the growing season. The reproductive phenology phases of each species were estimated through fitting the phenological scores to the Richards function. Important findings: Under soil water stress caused by warming, most plants in the alpine meadow advanced or delayed their reproductive events. As a result, warming significantly delayed phenological development of K. pygmaea. Warming significantly advanced reproductive phenology of P. saundersiana, S. purpurea and F. coelestis, but not of P. cuneata and Y. simulatrix. In addition, warming significantly shortened the average flowering duration of alpine plant species. The potentially warmer and drier growing seasons under climate change may shift the reproductive phenology of the alpine systems in similar pattern.     

The influence of life history traits on the phenological response of British butterflies to climate variability since the late‐19th century

Many species of plants and animals have advanced their phenology in response to climate warming in recent decades. Most of the evidence available for these shifts is based on data from the last few decades, a period coinciding with rapid climate warming. Baseline data is required to put these recent phenological changes in a long‐term context. We analysed the phenological response of 51 resident British butterfly species using data from 83 500 specimens in the collections of the Natural History Museum, London, covering the period 1880–1970. Our analysis shows that only three species significantly advanced their phenology between 1880 and 1970, probably reflecting the relatively small increase in spring temperature over this period. However, the phenology of all but one of the species we analysed showed phenological sensitivity to inter‐annual climate variability and a significant advancement in phenology in years in which spring or summer temperatures were warm and dry. The phenologies of butterfly species were more sensitive to weather if the butterfly species was early flying, southerly distributed, and a generalist in terms of larval diet. This observation is consistent with the hypothesis that species with greater niche breadth may be more phenologically sensitive than species with important niche constraints. Comparison of our results with post‐1976 data from the UK Butterfly Monitoring Scheme show that species flying early in the year had a greater rate of phenological advancement prior to the mid‐1970s. Additionally, prior to the mid‐1970s, phenology was influenced by temperatures in March or April, whereas since 1976, February temperature had a stronger influence on the phenology. These results suggest that early flying species may be approaching the limits of phenological advancement in response to recent climate warming.     

Effects of experimental warming on plant reproductive phenology in Xizang alpine meadow

Aims Climate warming strongly influences reproductive phenology of plants in alpine and arctic ecosystems. Here we focus on phenological shifts caused by warming in a typical alpine meadow on the Qinghai-Xizang Plateau. Our objective was to explore phenological responses of alpine plant species to experimental warming. Methods Passive warming was achieved using open-top chambers (OTCs). The treatments included control (C), and four levels of warming (T1, T2, T3, T4). We selected Kobresia pygmaea, Potentilla saundersiana, Potentilla cuneata, Stipa purpurea, Festuca coelestis and Youngia simulatrix as the focal species. Plant phenology was scored every 3-5 days in the growing season. The reproductive phenology phases of each species were estimated through fitting the phenological scores to the Richards function. Important findings Under soil water stress caused by warming, most plants in the alpine meadow advanced or delayed their reproductive events. As a result, warming significantly delayed phenological development of K. pygmaea. Warming significantly advanced reproductive phenology of P. saundersiana, S. purpurea and F. coelestis, but not of P. cuneata and Y. simulatrix. In addition, warming significantly shortened the average flowering duration of alpine plant species. The potentially warmer and drier growing seasons under climate change may shift the reproductive phenology of the alpine systems in similar pattern.     

实验增温对藏北高寒草甸植物繁殖物候的影响

全球气候变暖对高寒和极地地区的植物物候产生强烈的影响。该研究主要关注增温条件下藏北高寒草甸不同功能型植物繁殖时间(生殖物候)的改变。实验采用开顶箱式增温方法, 对3个主要功能群浅根-早花、浅根-中花和深根-晚花植物的现蕾、开花、结实时间进行观测。研究结果表明: (1)增温导致了土壤水分胁迫, 显著推迟了浅根-早花植物高山嵩草(Kobresia pygmaea)的繁殖时间; (2)增温显著提前了浅根-中花植物钉柱委陵菜(Potentilla saundersiana)和深根晚花植物紫花针茅(Stipa purpurea)和矮羊茅(Festuca coelestis)的繁殖时间; (3)增温没有显著影响浅根-中花植物楔叶委陵菜(Potentilla cuneata)和深根-晚花植物无茎黄鹌菜(Youngia simulatrix)的繁殖时间; (4)增温缩短了3种类型植物的开花持续时间。这些结果显示增温改变了藏北高寒草甸群落中多数物种的繁殖时间, 这预示着在未来更热更干的生长季, 青藏高原高寒草甸系统的植物物候格局可能会被重塑。     

Advancing breeding phenology in response to environmental change in a wild red deer population

Most evidence for advances in phenology of in response to recent climate warming in wild vertebrate populations has come from long‐term studies of birds. Few studies have either documented phenological advances or tested their climatic causes and demographic consequences in wild mammal systems. Using a long‐term study of red deer on the Isle of Rum, Scotland, we present evidence of significant temporal trends in six phenological traits: oestrus date and parturition date in females, and antler cast date, antler clean date, rut start date and rut end date in males. These traits advanced by between 5 and 12 days across a 28‐year study period. Local climate measures associated with plant growth in spring and summer (growing degree days) increased significantly over time and explained a significant amount of variation in all six phenological traits, largely accounting for temporal advances observed in some of the traits. However, there was no evidence for temporal changes in key female reproductive performance traits (offspring birth weight and offspring survival) in this population, despite significant relationships between these traits and female phenology. In males, average antler weights increased over time presumably as a result of improved resource availability and physiological condition through spring and summer. There was no evidence for any temporal change in average male annual breeding success, as might be expected if the timing of male rutting behaviour was failing to track advances in the timing of oestrus in females. Our results provide rare evidence linking phenological advances to climate warming in a wild mammal and highlight the potential complexity of relationships between climate warming, phenology and demography in wild vertebrates.     

Different temperature perception in high‐elevation plants: new insight into phenological development and implications for climate change in the alpine tundra

In alpine habitats, predicted warmer and longer growing seasons will influence plant phenology, with important implications for species adaptation and vegetation dynamics. However, little is known on the temperature sensitivity of different phenophases and on the characteristics allowing phenological variation among and within species. By integrating interannual micro‐climatic variability with experimental warming, we explored how the phenology of three alpine species is influenced by temperature and what mechanisms underlie intra‐ and inter‐specific phenological differences. The present study demonstrated that alpine plants have different temperature responses during their reproductive cycle, do not have constant thermal thresholds and heat‐use efficiencies to achieve the seed dispersal stage and can change their temperature sensitivity to flower along snowmelt gradients. In addition, the length of the reproductive cycle, which proved to be species‐specific under experimental warming, does not seem to be the only life‐history trait under selective pressure due to the short‐length of the snow‐free period. In a warming climate scenario, the phenology of sexual reproduction will be considerably altered, and alpine plants may be subjected to changes in population dynamics driven by altered perception of environmental cues appropriate for coordinating the timing of key life‐history events.     

Prairie plant phenology driven more by temperature than moisture in climate manipulations across a latitudinal gradient in the Pacific Northwest,USA

Plant phenology will likely shift with climate change, but how temperature and/or moisture regimes will control phenological responses is not well understood. This is particularly true in Mediterranean climate ecosystems where the warmest temperatures and greatest moisture availability are seasonally asynchronous. We examined plant phenological responses at both the population and community levels to four climate treatments (control, warming, drought, and warming plus additional precipitation) embedded within three prairies across a 520 km latitudinal Mediterranean climate gradient within the Pacific Northwest, USA. At the population level, we monitored flowering and abundances in spring 2017 of eight range‐restricted focal species planted both within and north of their current ranges. At the community level, we used normalized difference vegetation index (NDVI) measured from fall 2016 to summer 2018 to estimate peak live biomass, senescence, seasonal patterns, and growing season length. We found that warming exerted a stronger control than our moisture manipulations on phenology at both the population and community levels. Warming advanced flowering regardless of whether a species was within or beyond its current range. Importantly, many of our focal species had low abundances, particularly in the south, suggesting that establishment, in addition to phenological shifts, may be a strong constraint on their future viability. At the community level, warming advanced the date of peak biomass regardless of site or year. The date of senescence advanced regardless of year for the southern and central sites but only in 2018 for the northern site. Growing season length contracted due to warming at the southern and central sites (~3 weeks) but was unaffected at the northern site. Our results emphasize that future temperature changes may exert strong influence on the timing of a variety of plant phenological events, especially those events that occur when temperature is most limiting, even in seasonally water‐limited Mediterranean ecosystems.     

Fractional coverage rather than green chromatic coordinate is a robust indicator to track grassland phenology using smartphone photography

Cost-effective and accurate methods are critical in monitoring grassland phenology. Smartphones have great potential for tracking phenological changes owing to their easy operation and flexible sampling, especially for experimental treatments on grassland phenology at local scale. However, it remains an open question for an accurate and robust measure in tracking plant phenology under various light conditions when using smartphones. To fill this gap, we compared green chromatic coordinate (GCC) and fractional coverage (FC) produced by three widely-used smartphone types across varying light conditions. We also explored the effects of experimental warming and N addition on grassland spring phenology using smartphone photography. We found that light conditions affected GCC values derived from various smartphones. The percentile method did not reduce the influence of various light conditions on GCC calculation. When using one type of smartphone, other techniques (e.g., shading strong sunlight) were required to minimize the effect of light on mean GCC during photography, but they did not work for FC. Based on this method, three smartphone types generated similar phenological metrics (i.e., start, peak and end of growing season) detected from GCC or FC. GCC and FC produced synchronous seasonal trajectories and similar phenological metrics throughout the growing season. We also found that experimental warming advanced spring phenology in the relatively wet year of 2018, but delayed it in the dry year of 2019. In contrast, N addition had little influence on spring phenology. Our results revealed that overlooking the influence of ambient light and smartphone types on GCC might lead to uncertainties in phenological measurement. Robust FC metrics extracted from smartphone photography detected grassland phenology differences in response to warming and N deposition. These findings provide evidence that FC is a robust indicator to accurately monitor grassland phenology using smartphone photography. Their utility resonates with those monitoring many experimental plots or sites with various light conditions.     

Detecting nonlinear response of spring phenology to climate change by Bayesian analysis

The impact of climate change on the advancement of plant phenological events has been heavily studied in the last decade. Although the majority of spring plant phenological events have been trending earlier, this is not universally true. Recent work has suggested that species that are not advancing in their spring phenological behavior are responding more to lack of winter chill than increased spring heat. One way to test this hypothesis is by evaluating the behavior of a species known to have a moderate to high chilling requirement and examining how it is responding to increased warming. This study used a 60‐year data set for timing of leaf‐out and male flowering of walnut (Juglans regia) cultivar ‘Payne’ to examine this issue. The spring phenological behavior of ‘Payne’ walnut differed depending on bud type. The vegetative buds, which have a higher chilling requirement, trended toward earlier leaf‐out until about 1994, when they shifted to later leaf‐out. The date of male bud pollen shedding advanced over the course of the whole record. Our findings suggest that many species which have exhibited earlier bud break are responding to warmer spring temperatures, but may shift into responding more to winter temperatures (lack of adequate chilling) as warming continues.     

基于OTCs模拟增温方式探讨气候变暖对青藏高原草地生态系统的影响

开顶式生长室(OTCs)增温实验是研究全球气候变化与陆地生态系统关系的主要方法之一,已广泛应用于青藏高原地区。该文通过对近些年国内外研究文献的回顾,分别从植物物候、群落结构、生物量和土壤方面综合分析青藏高原草地生态系统对OTCs模拟增温实验的响应。研究发现:增温使群落返青期提前、枯黄期延迟,生长季延长;有利于禾本科植物的生长;高寒草甸地下生物量分配格局向深层转移;高寒草地生态系统对模拟增温的响应存在不确定性,受到地域、群落类型和实验时间的影响;在增温条件下,降雨和冻土融化引起的土壤水分变化通过调控生态系统的物候、生产力、土壤等途径控制着生态系统对气候变暖的响应。并在此基础上,提出了将来应着重研究的几个方面。     

Drought advances spring growth phenology of the Mediterranean shrub Erica multiflora

Current climate projections predict drier and warmer conditions in the Mediterranean basin over the next century. While advanced spring growth due to warming has been described in the literature, few data are available on the effects of drought on phenology. Hence, the phenology and growth of two Mediterranean shrubs, Erica multiflora and Globularia alypum, was studied in a rainfall exclusion field experiment to simulate spring drought in a natural shrubland. We estimated the onset of growth in spring by monitoring the appearance of new stems, and the end of growth in summer by following the elongation of stems. Drought treatment caused earlier onset of the spring growing season in E. multiflora, whereas no advance was observed in G. alypum. However, growth cessation was not affected in E. multiflora. Drought reduced the growth of both shrubs, as reflected in less stem elongation. The results show that a drier climate might affect not only growth but also spring phenology of some Mediterranean species. We suggest that a reduction in the cooling effect of transpiration may have analogous effects to warming and might advance the start of growth in E. multiflora, a species whose phenology has been described as warming‐sensitive. The lengthening of the growing season resulting from advanced growth did not imply higher productivity, as growth was restricted by drought.     

内蒙古中西部优势植物春季返青对降水的非线性响应

长期以来,研究植被物候变化与气候因子的关系多基于线性模型,事实上植被物候对气候变化的响应可能是非线性的。以1984-2017年内蒙古中西部温性典型草原和温性草原化荒漠长时间序列植被物候观测资料为基础,分析了近40年来气候变化背景下不同草地类型优势植物返青期变化特征及其对春季降水的非线性响应。结果表明：（1）温性典型草原冷蒿返青期主要受水分控制,与春季降水量表现为开口向下的二次函数关系。气候暖干化导致的水分亏缺是冷蒿返青期呈极显著延迟趋势（1.32 d/a）的主要原因;春季降水量超过60 mm时,冷蒿返青期表现出由延迟转变为提前的趋势。（2）温性草原化荒漠猫头刺返青期主要受热量控制。受春季显著升温影响,猫头刺返青期呈极显著提前趋势（0.63 d/a）。春季降水增多利于猫头刺提早返青,二者表现为开口向上的二次函数关系;春季降水量超过40 mm时,猫头刺返青期对降水的响应程度逐渐降低,这可能与荒漠植物本身的生理特性有关。     

A Range-Expanding Shrub Species Alters Plant Phenological Response to Experimental Warming

Shifts in plant species phenology (the timing of life-history events such as flowering) have been observed worldwide in concert with rising global temperatures. While most species display earlier phenology with warming, there is large variation among, and even within, species in phenological sensitivity to rising temperatures. Other indirect effects of climate change, such as shifting species composition and altered species interactions, may also be contributing to shifting plant phenology. Here, we describe how experimental warming and the presence of a range-expanding species, sagebrush (Artemisia rothrockii), interact to influence the flowering phenology (day of first and peak flowering) and production (number of flowers) of an alpine cushion plant, Trifolium andersonii, in California’s White Mountains. Both first flowering and peak flowering were strongly accelerated by warming, but not when sagebrush was present. Warming significantly increased flower production of T. andersonii, but less so in the presence of sagebrush. A shading treatment delayed phenology and lowered flower production, suggesting that shading may be the mechanism by which sagebrush presence delayed flowering of the understory species. This study demonstrates that species interactions can modify phenological responses to climate change, and suggests that indirect effects of rising temperatures arising from shifting species ranges and altered species interactions may even exceed the direct effects of rising temperatures on phenology.     

Eco‐evolutionary responses of Bromus tectorum to climate change: implications for biological invasions

How plant populations, communities, and ecosystems respond to climate change is a critical focus in ecology today. The responses of introduced species may be especially rapid. Current models that incorporate temperature and precipitation suggest that future Bromus tectorum invasion risk is low for the Colorado Plateau. With a field warming experiment at two sites in southeastern Utah, we tested this prediction over 4 years, measuring B. tectorum phenology, biomass, and reproduction. In a complimentary greenhouse study, we assessed whether changes in field B. tectorum biomass and reproductive output influence offspring performance. We found that following a wet winter and early spring, the timing of spring growth initiation, flowering, and summer senescence all advanced in warmed plots at both field sites and the shift in phenology was progressively larger with greater warming. Earlier green‐up and development was associated with increases in B. tectorum biomass and reproductive output, likely due early spring growth, when soil moisture was not limiting, and a lengthened growing season. Seeds collected from plants grown in warmed plots had higher biomass and germination rates and lower mortality than seeds from ambient plots. However, in the following two dry years, we observed no differences in phenology between warmed and ambient plots. In addition, warming had a generally negative effect on B. tectorum biomass and reproduction in dry years and this negative effect was significant in the plots that received the highest warming treatment. In contrast to models that predict negative responses of B. tectorum to warmer climate on the Colorado Plateau, the effects of warming were more nuanced, relied on background climate, and differed between the two field sites. Our results highlight the importance of considering the interacting effects of temperature, precipitation, and site‐specific characteristics such as soil texture, on plant demography and have direct implications for B. tectorum invasion dynamics on the Colorado Plateau.     

Flowering phenology and reproductive fitness along a mountain slope: maladaptive responses to transplantation to a warmer climate in Campanula thyrsoides

In many biomes, global warming has resulted in advanced and longer growing seasons, which has often led to earlier flowering in plant taxa. Elevational gradients are ideal to study the effects of global warming as they allow transplantation of plants from their original cooler higher elevations down to elevations with a prospective climate. We transplanted plants from ten populations of the European alpine monocarpic herb species Campanula thyrsoides L. to three sites along a steep mountain slope (600, 1,235 and 1,850 m above sea level) in the Swiss Alps and asked whether reproductive phenology adjusts plastically to elevation and if these responses were adaptive, i.e. increased the fitness of plants. We further assessed current genetic differentiation in phenotypic traits and whether any such origin effects were due to adaptation to climatic conditions of origin. Our results showed that transplantation to lower elevations caused strong shifts in phenology, with plants starting growth and flowering earlier than plants placed at higher elevations. However, compared to flower production at high elevation, number of flowers per plant decreased 21 % at mid- and 61 % at low elevation. The shift in phenology thus came with a high cost in fitness, and we suggest that phenology is maladaptive when C. thyrsoides faces temperature conditions deviating from its natural amplitude. We conclude that the frequently reported phenological shift in plant species as a response to global warming may include heavy fitness costs that may hamper species survival.     

Shifts in the temperature‐sensitive periods for spring phenology in European beech and pedunculate oak clones across latitudes and over recent decades

Spring phenology of temperate trees has advanced worldwide in response to global warming. However, increasing temperatures may not necessarily lead to further phenological advance, especially in the warmer latitudes because of insufficient chilling and/or shorter day length. Determining the start of the forcing phase, that is, when buds are able to respond to warmer temperatures in spring, is therefore crucial to predict how phenology will change in the future. In this study, we used 4,056 leaf‐out date observations during the period 1969–2017 for clones of European beech (Fagus sylvatica L.) and pedunculate oak (Quercus robur L.) planted in 63 sites covering a large latitudinal gradient (from Portugal ~41°N to Norway ~63°N) at the International Phenological Gardens in order to (a) evaluate how the sensitivity periods to forcing and chilling have changed with climate warming, and (b) test whether consistent patterns occur along biogeographical gradients, that is, from colder to warmer environments. Partial least squares regressions suggest that the length of the forcing period has been extended over the recent decades with climate warming in the colder latitudes but has been shortened in the warmer latitudes for both species, with a more pronounced shift for beech. We attribute the lengthening of the forcing period in the colder latitudes to earlier opportunities with temperatures that can promote bud development. In contrast, at warmer or oceanic climates, the beginning of the forcing period has been delayed, possibly due to insufficient chilling. However, in spite of a later beginning of the forcing period, spring phenology has continued to advance at these areas due to a faster satisfaction of heat requirements induced by climate warming. Overall, our results support that ongoing climate warming will have different effects on the spring phenology of forest trees across latitudes due to the interactions between chilling, forcing and photoperiod.