Unexpected role of winter precipitation in determining heat requirement for spring vegetation green‐up at northern middle and high latitudes

Heat requirement, expressed in growing degree days (GDD), is a widely used method to assess and predict the effect of temperature on plant development. Until recently, the analysis of spatial patterns of GDD requirement for spring vegetation green‐up onset was limited to local and regional scales, mainly because of the sparse and aggregated spatial availability of ground phenology data. Taking advantage of the large temporal and spatial scales of remote sensing‐based green‐up onset data, we studied the spatial patterns of GDD requirement for vegetation green‐up at northern middle and high latitudes. We further explored the correlations between GDD requirement for vegetation green‐up and previous winter season chilling temperatures and precipitation, using spatial partial correlations. We showed that GDD requirement for vegetation green‐up onset declines towards the north at a mean rate of 18.8 °C‐days per degree latitude between 35°N and 70°N, and vary significantly among different vegetation types. Our results confirmed that the GDD requirement for vegetation green‐up is negatively correlated with previous winter chilling, which was defined as the number of chilling days from the day when the land surface froze in the previous autumn to the day of green‐up onset. This negative correlation is a well‐known phenomenon from local studies. Interestingly, irrespective of the vegetation type, we also found a positive correlation between the GDD requirement and previous winter season precipitation, which was defined as the sum of the precipitation of the month when green‐up onset occur and the precipitation that occurred during the previous 2 months. Our study suggests that GDD requirement, chilling and precipitation may have complex interactions in their effects on spring vegetation green‐up phenology. These findings have important implications for improving phenology models and could therefore advance our understanding of the interplay between spring phenology and carbon fluxes.     

Chilling and forcing temperatures interact to predict the onset of wood formation in Northern Hemisphere conifers

The phenology of wood formation is a critical process to consider for predicting how trees from the temperate and boreal zones may react to climate change. Compared to leaf phenology, however, the determinism of wood phenology is still poorly known. Here, we compared for the first time three alternative ecophysiological model classes (threshold models, heat‐sum models and chilling‐influenced heat‐sum models) and an empirical model in their ability to predict the starting date of xylem cell enlargement in spring, for four major Northern Hemisphere conifers (Larix decidua, Pinus sylvestris, Picea abies and Picea mariana). We fitted models with Bayesian inference to wood phenological data collected for 220 site‐years over Europe and Canada. The chilling‐influenced heat‐sum model received most support for all the four studied species, predicting validation data with a 7.7‐day error, which is within one day of the observed data resolution. We conclude that both chilling and forcing temperatures determine the onset of wood formation in Northern Hemisphere conifers. Importantly, the chilling‐influenced heat‐sum model showed virtually no spatial bias whichever the species, despite the large environmental gradients considered. This suggests that the spring onset of wood formation is far less affected by local adaptation than by environmentally driven plasticity. In a context of climate change, we therefore expect rising winter–spring temperature to exert ambivalent effects on the spring onset of wood formation, tending to hasten it through the accumulation of forcing temperature, but imposing a higher forcing temperature requirement through the lower accumulation of chilling.     

Increased heat requirement for leaf flushing in temperate woody species over 1980–2012: effects of chilling,precipitation and insolation

Recent studies have revealed large unexplained variation in heat requirement‐based phenology models, resulting in large uncertainty when predicting ecosystem carbon and water balance responses to climate variability. Improving our understanding of the heat requirement for spring phenology is thus urgently needed. In this study, we estimated the species‐specific heat requirement for leaf flushing of 13 temperate woody species using long‐term phenological observations from Europe and North America. The species were defined as early and late flushing species according to the mean date of leaf flushing across all sites. Partial correlation analyses were applied to determine the temporal correlations between heat requirement and chilling accumulation, precipitation and insolation sum during dormancy. We found that the heat requirement for leaf flushing increased by almost 50% over the study period 1980–2012, with an average of 30 heat units per decade. This temporal increase in heat requirement was observed in all species, but was much larger for late than for early flushing species. Consistent with previous studies, we found that the heat requirement negatively correlates with chilling accumulation. Interestingly, after removing the variation induced by chilling accumulation, a predominantly positive partial correlation exists between heat requirement and precipitation sum, and a predominantly negative correlation between heat requirement and insolation sum. This suggests that besides the well‐known effect of chilling, the heat requirement for leaf flushing is also influenced by precipitation and insolation sum during dormancy. However, we hypothesize that the observed precipitation and insolation effects might be artefacts attributable to the inappropriate use of air temperature in the heat requirement quantification. Rather than air temperature, meristem temperature is probably the prominent driver of the leaf flushing process, but these data are not available. Further experimental research is thus needed to verify whether insolation and precipitation sums directly affect the heat requirement for leaf flushing.     

Predicting a change in the order of spring phenology in temperate forests

The rise in spring temperatures over the past half‐century has led to advances in the phenology of many nontropical plants and animals. As species and populations differ in their phenological responses to temperature, an increase in temperatures has the potential to alter timing‐dependent species interactions. One species‐interaction that may be affected is the competition for light in deciduous forests, where early vernal species have a narrow window of opportunity for growth before late spring species cast shade. Here we consider the Marsham phenology time series of first leafing dates of thirteen tree species and flowering dates of one ground flora species, which spans two centuries. The exceptional length of this time series permits a rare comparison of the statistical support for parameter‐rich regression and mechanistic thermal sensitivity phenology models. While mechanistic models perform best in the majority of cases, both they and the regression models provide remarkably consistent insights into the relative sensitivity of each species to forcing and chilling effects. All species are sensitive to spring forcing, but we also find that vernal and northern European species are responsive to cold temperatures in the previous autumn. Whether this sensitivity reflects a chilling requirement or a delaying of dormancy remains to be tested. We then apply the models to projected future temperature data under a fossil fuel intensive emissions scenario and predict that while some species will advance substantially others will advance by less and may even be delayed due to a rise in autumn and winter temperatures. Considering the projected responses of all fourteen species, we anticipate a change in the order of spring events, which may lead to changes in competitive advantage for light with potential implications for the composition of temperate forests.     

Simulating the onset of spring vegetation growth across the Northern Hemisphere

Changes in the spring onset of vegetation growth in response to climate change can profoundly impact climate–biosphere interactions. Thus, robust simulation of spring onset is essential to accurately predict ecosystem responses and feedback to ongoing climate change. To date, the ability of vegetation phenology models to reproduce spatiotemporal patterns of spring onset at larger scales has not been thoroughly investigated. In this study, we took advantage of phenology observations via remote sensing to calibrate and evaluated six models, including both one‐phase (considering only forcing temperatures) and two‐phase (involving forcing, chilling, and photoperiod) models across the Northern Hemisphere between 1982 and 2012. Overall, we found that the model that integrated the photoperiod effect performed best at capturing spatiotemporal patterns of spring phenology in boreal and temperate forests. By contrast, all of the models performed poorly in simulating the onset of growth in grasslands. These results suggest that the photoperiod plays a role in controlling the onset of growth in most Northern Hemisphere forests, whereas other environmental factors (e.g., precipitation) should be considered when simulating the onset of growth in grasslands. We also found that the one‐phase model performed as well as the two‐phase models in boreal forests, which implies that the chilling requirement is probably fulfilled across most of the boreal zone. Conversely, two‐phase models performed better in temperate forests than the one‐phase model, suggesting that photoperiod and chilling play important roles in these temperate forests. Our results highlight the significance of including chilling and photoperiod effects in models of the spring onset of forest growth at large scales, and indicate that the consideration of additional drivers may be required for grasslands.     

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.     

Chilling outweighs photoperiod in preventing precocious spring development

It is well known that increased spring temperatures cause earlier onset dates of leaf unfolding and flowering. However, a temperature increase in winter may be associated with delayed development when species' chilling requirements are not fulfilled. Furthermore, photosensitivity is supposed to interfere with temperature triggers. To date, neither the relative importance nor possible interactions of these three factors have been elucidated. In this study, we present a multispecies climate chamber experiment to test the effects of chilling and photoperiod on the spring phenology of 36 woody species. Several hypotheses regarding their variation with species traits (successional strategy, floristic status, climate of their native range) were tested. Long photoperiods advanced budburst for one‐third of the studied species, but magnitudes of these effects were generally minor. In contrast to prior hypotheses, photosensitive responses were not restricted to climax or oceanic species. Increased chilling length advanced budburst for almost all species; its effect greatly exceeding that of photoperiod. Moreover, we suggest that photosensitivity and chilling effects have to be rigorously disentangled, as the response to photoperiod was restricted to individuals that had not been fully chilled. The results indicate that temperature requirements and successional strategy are linked, with climax species having higher chilling and forcing requirements than pioneer species. Temperature requirements of invasive species closely matched those of native species, suggesting that high phenological concordance is a prerequisite for successful establishment. Lack of chilling not only led to a considerable delay in budburst but also caused substantial changes in the chronological order of species' budburst. The results reveal that increased winter temperatures might impact forest ecosystems more than formerly assumed. Species with lower chilling requirements, such as pioneer or invasive species, might profit from warming winters, if late spring frost events would in parallel occur earlier.     

From observations to experiments in phenology research: investigating climate change impacts on trees and shrubs using dormant twigs

Background and Aims Climate change is advancing the leaf-out times of many plant species and mostly extending the growing season in temperate ecosystems. Laboratory experiments using twig cuttings from woody plant species present an affordable, easily replicated approach to investigate the relative importance of factors such as winter chilling, photoperiod, spring warming and frost tolerance on the leafing-out times of plant communities. This Viewpoint article demonstrates how the results of these experiments deepen our understanding beyond what is possible via analyses of remote sensing and field observation data, and can be used to improve climate change forecasts of shifts in phenology, ecosystem processes and ecological interactions.Scope The twig method involves cutting dormant twigs from trees, shrubs and vines on a single date or at intervals over the course of the winter and early spring, placing them in containers of water in controlled environments, and regularly recording leaf-out, flowering or other phenomena. Prior to or following leaf-out or flowering, twigs may be assigned to treatment groups for experiments involving temperature, photoperiod, frost, humidity and more. Recent studies using these methods have shown that winter chilling requirements and spring warming strongly affect leaf-out and flowering times of temperate trees and shrubs, whereas photoperiod requirements are less important than previously thought for most species. Invasive plant species have weaker winter chilling requirements than native species in temperate ecosystems, and species that leaf-out early in the season have greater frost tolerance than later leafing species.Conclusions This methodology could be extended to investigate additional drivers of leaf-out phenology, leaf senescence in the autumn, and other phenomena, and could be a useful tool for education and outreach. Additional ecosystems, such as boreal, southern hemisphere and sub-tropical forests, could also be investigated using dormant twigs to determine the drivers of leaf-out times and how these ecosystems will be affected by climate change.     

Temperature alone does not explain phenological variation of diverse temperate plants under experimental warming

Anthropogenic climate change has altered temperate forest phenology, but how these trends will play out in the future is controversial. We measured the effect of experimental warming of 0.6–5.0 °C on the phenology of a diverse suite of 11 plant species in the deciduous forest understory (Duke Forest, North Carolina, USA) in a relatively warm year (2011) and a colder year (2013). Our primary goal was to dissect how temperature affects timing of spring budburst, flowering, and autumn leaf coloring for functional groups with different growth habits, phenological niches, and xylem anatomy. Warming advanced budburst of six deciduous woody species by 5–15 days and delayed leaf coloring by 18–21 days, resulting in an extension of the growing season by as much as 20–29 days. Spring temperature accumulation was strongly correlated with budburst date, but temperature alone cannot explain the diverse budburst responses observed among plant functional types. Ring‐porous trees showed a consistent temperature response pattern across years, suggesting these species are sensitive to photoperiod. Conversely, diffuse‐porous species responded differently between years, suggesting winter chilling may be more important in regulating budburst. Budburst of the ring‐porous Quercus alba responded nonlinearly to warming, suggesting evolutionary constraints may limit changes in phenology, and therefore productivity, in the future. Warming caused a divergence in flowering times among species in the forest community, resulting in a longer flowering season by 10‐16 days. Temperature was a good predictor of flowering for only four of the seven species studied here. Observations of interannual temperature variability overpredicted flowering responses in spring‐blooming species, relative to our warming experiment, and did not consistently predict even the direction of flowering shifts. Experiments that push temperatures beyond historic variation are indispensable for improving predictions of future changes in phenology.     

Chilling and heat requirements for leaf unfolding in European beech and sessile oak populations at the southern limit of their distribution range

With global warming, an advance in spring leaf phenology has been reported worldwide. However, it is difficult to forecast phenology for a given species, due to a lack of knowledge about chilling requirements. We quantified chilling and heat requirements for leaf unfolding in two European tree species and investigated their relative contributions to phenological variations between and within populations. We used an extensive database containing information about the leaf phenology of 14 oak and 10 beech populations monitored over elevation gradients since 2005. In parallel, we studied the various bud dormancy phases, in controlled conditions, by regularly sampling low- and high-elevation populations during fall and winter. Oak was 2.3 times more sensitive to temperature for leaf unfolding over the elevation gradient and had a lower chilling requirement for dormancy release than beech. We found that chilling is currently insufficient for the full release of dormancy, for both species, at the lowest elevations in the area studied. Genetic variation in leaf unfolding timing between and within oak populations was probably due to differences in heat requirement rather than differences in chilling requirement. Our results demonstrate the importance of chilling for leaf unfolding in forest trees and indicate that the advance in leaf unfolding phenology with increasing temperature will probably be less pronounced than forecasted. This highlights the urgent need to determine experimentally the interactions between chilling and heat requirements in forest tree species, to improve our understanding and modeling of changes in phenological timing under global warming.     

Chilling rather than photoperiod controls budburst for gymnosperm species in subtropical China

低温而不是光周期调控中国亚热带裸子植物的出芽物候 摘要：被子植物春季物候的调控机制已经得到了广泛的研究。然而,裸子植物和被子植物在3亿年前就产生分化,裸子植物与被子植物的物候可能是受不同的因素所调控。亚热带植物物候的调节机制在很大程度上尚不明确,亚热带裸子植物物候是否由冷激需求和光照调控仍未得到验证。本研究在人工气候箱中设置了3个冷激处理和3 个光周期处理,并对切枝的出芽期进行了为期8周的研究。实验中我们测试了8种裸子植物：柳杉(Cryptomeria japonica)、杉木(Cunninghamia lanceolata)、柏树(Cupressus funebris)、银杏(Ginkgo biloba)、水杉(Metasequoia glyptostroboides)、马尾松(Pinus massoniana)、金钱 松(Pseudolarix amabilis)和罗汉松(Podocarpus macrophyllus),检验其出芽物候是否对光周期敏感或者是否具有较强的冷激需求,以及这两种因素哪个对促进出芽更为重要。研究结果表明,对于裸子植物,冷 激促进了出芽并提高了出芽率,而且裸子植物需要适度的低温天数来实现出芽。有趣的是,在同一森 林中裸子植物比被子植物对积温的需求更高。与德国温带裸子植物(194–600 d · °C)相比,亚热带裸子植 物(814–1150 d · °C)对积温的需求更高。光周期对裸子植物出芽的影响较小,说明冷激对裸子植物出芽的 促进作用大于光周期。这些结果表明,随着全球气候持续变暖,冬季气温的升高不仅会影响亚热带被子植物也会影响裸子植物的物候,从而可能导致春季出芽期的延迟。     

Experimental warming alters spring phenology of certain plant functional groups in an early successional forest community

Experimental study of the effects of projected climate change on plant phenology allows us to isolate effects of warming on life‐history events such as leaf out. We simulated a 2 °C temperature increase and 20% precipitation increase in a recently harvested temperate deciduous forest community in central Pennsylvania, USA, and observed the leaf out phenology of all species in 2009 and 2010. Over 130 plant species were monitored weekly in study plots, but due to high variability in species composition among plots, species were grouped into five functional groups: short forbs, tall forbs, shrubs, small trees, and large trees. Tall forbs and large trees, which usually emerge in the late spring, advanced leaf out 14–18 days in response to warming. Short forbs, shrubs, and small trees emerge early in spring and did not alter their phenology in response to warming or increased precipitation treatments. Earlier leaf out of tall forbs and large trees coincided with almost 3 weeks of increased community‐level leaf area index, indicating greater competition and a condensed spring green‐up period. While phenology of large trees and tall forbs appears to be strongly influenced by temperature‐based growth cues, our results suggest that photoperiod and chilling cues more strongly influence the leaf out of other functional groups. Reduced freeze events and warmer temperatures from predicted climate change will interact with nontemperature growth cues to have cascading consequences throughout the ecosystem.     

Changes in the structure and function of northern Alaskan ecosystems when considering variable leaf‐out times across groupings of species in a dynamic vegetation model

The phenology of arctic ecosystems is driven primarily by abiotic forces, with temperature acting as the main determinant of growing season onset and leaf budburst in the spring. However, while the plant species in arctic ecosystems require differing amounts of accumulated heat for leaf‐out, dynamic vegetation models simulated over regional to global scales typically assume some average leaf‐out for all of the species within an ecosystem. Here, we make use of air temperature records and observations of spring leaf phenology collected across dominant groupings of species (dwarf birch shrubs, willow shrubs, other deciduous shrubs, grasses, sedges, and forbs) in arctic and boreal ecosystems in Alaska. We then parameterize a dynamic vegetation model based on these data for four types of tundra ecosystems (heath tundra, shrub tundra, wet sedge tundra, and tussock tundra), as well as ecotonal boreal white spruce forest, and perform model simulations for the years 1970–2100. Over the course of the model simulations, we found changes in ecosystem composition under this new phenology algorithm compared with simulations with the previous phenology algorithm. These changes were the result of the differential timing of leaf‐out, as well as the ability for the groupings of species to compete for nitrogen and light availability. Regionally, there were differences in the trends of the carbon pools and fluxes between the new phenology algorithm and the previous phenology algorithm, although these differences depended on the future climate scenario. These findings indicate the importance of leaf phenology data collection by species and across the various ecosystem types within the highly heterogeneous Arctic landscape, and that dynamic vegetation models should consider variation in leaf‐out by groupings of species within these ecosystems to make more accurate projections of future plant distributions and carbon cycling in Arctic regions.     

Long‐term changes in the impacts of global warming on leaf phenology of four temperate tree species

Contrary to the generally advanced spring leaf unfolding under global warming, the effects of the climate warming on autumn leaf senescence are highly variable with advanced, delayed, and unchanged patterns being all reported. Using one million records of leaf phenology from four dominant temperate species in Europe, we investigated the temperature sensitivities of spring leaf unfolding and autumn leaf senescence (S_T, advanced or delayed days per degree Celsius). The S_T of spring phenology in all of the four examined species showed an increase and decrease during 1951–1980 and 1981–2013, respectively. The decrease in the S_T during 1981–2013 appears to be caused by reduced accumulation of chilling units. As with spring phenology, the S_T of leaf senescence of early successional and exotic species started to decline since 1980. In contrast, for late successional species, the S_T of autumn senescence showed an increase for the entire study period from 1951 to 2013. Moreover, the impacts of rising temperature associated with global warming on spring leaf unfolding were stronger than those on autumn leaf senescence. The timing of leaf senescence was positively correlated with the timing of leaf unfolding during 1951–1980. However, as climate warming continued, the differences in the responses between spring and autumn phenology gradually increased, so that the correlation was no more significant during 1981–2013. Our results further suggest that since 2000, due to the decreased temperature sensitivity of leaf unfolding the length of the growing season has not increased any more. These finding needs to be addressed in vegetation models used for assessing the effects of climate change.     

Spring predictability explains different leaf‐out strategies in the woody floras of North America,Europe and East Asia

Intuitively, interannual spring temperature variability (STV) should influence the leaf‐out strategies of temperate zone woody species, with high winter chilling requirements in species from regions where spring warming varies greatly among years. We tested this hypothesis using experiments in 215 species and leaf‐out monitoring in 1585 species from East Asia (EA), Europe (EU) and North America (NA). The results reveal that species from regions with high STV indeed have higher winter chilling requirements, and, when grown under the same conditions, leaf out later than related species from regions with lower STV. Since 1900, STV has been consistently higher in NA than in EU and EA, and under experimentally short winter conditions NA species required 84% more spring warming for bud break, EU ones 49% and EA ones only 1%. These previously unknown continental‐scale differences in phenological strategies underscore the need for considering regional climate histories in global change models.     

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.     

Daylength helps temperate deciduous trees to leaf‐out at the optimal time

Global warming has led to substantially earlier spring leaf‐out in temperate‐zone deciduous trees. The interactive effects of temperature and daylength underlying this warming response remain unclear. However, they need to be accurately represented by earth system models to improve projections of the carbon and energy balances of temperate forests and the associated feedbacks to the Earth's climate system. We studied the control of leaf‐out by daylength and temperature using data from six tree species across 2,377 European phenological network ( www.pep725.eu ), each with at least 30 years of observations. We found that, in addition to and independent of the known effect of chilling, daylength correlates negatively with the heat requirement for leaf‐out in all studied species. In warm springs when leaf‐out is early, days are short and the heat requirement is higher than in an average spring, which mitigates the warming‐induced advancement of leaf‐out and protects the tree against precocious leaf‐out and the associated risks of late frosts. In contrast, longer‐than‐average daylength (in cold springs when leaf‐out is late) reduces the heat requirement for leaf‐out, ensuring that trees do not leaf‐out too late and miss out on large amounts of solar energy. These results provide the first large‐scale empirical evidence of a widespread daylength effect on the temperature sensitivity of leaf‐out phenology in temperate deciduous trees.     

Climate warming increases spring phenological differences among temperate trees

Climate warming has substantially advanced spring leaf flushing, but winter chilling and photoperiod co‐determine the leaf flushing process in ways that vary among species. As a result, the interspecific differences in spring phenology (IDSP) are expected to change with climate warming, which may, in turn, induce negative or positive ecological consequences. However, the temporal change of IDSP at large spatiotemporal scales remains unclear. In this study, we analyzed long‐term in‐situ observations (1951–2016) of six, coexisting temperate tree species from 305 sites across Central Europe and found that phenological ranking did not change when comparing the rapidly warming period 1984–2016 to the marginally warming period 1951–1983. However, the advance of leaf flushing was significantly larger in early‐flushing species EFS (6.7 ± 0.3 days) than in late‐flushing species LFS (5.9 ± 0.2 days) between the two periods, indicating extended IDSP. This IDSP extension could not be explained by differences in temperature sensitivity between EFS and LFS; however, climatic warming‐induced heat accumulation effects on leaf flushing, which were linked to a greater heat requirement and higher photoperiod sensitivity in LFS, drove the shifts in IDSP. Continued climate warming is expected to further extend IDSP across temperate trees, with associated implications for ecosystem function.     

Shifting and extension of phenological periods with increasing temperature along elevational transects in southern Bavaria

The impact of global warming on phenology has been widely studied, and almost consistently advancing spring events have been reported. Especially in alpine regions, an extraordinary rapid warming has been observed in the last decades. However, little is known about phenological phases over the whole vegetation period at high elevations. We observed 12 phenological phases of seven tree species and measured air temperature at 42 sites along four transects of about 1000 m elevational range in the years 2010 and 2011 near Garmisch‐Partenkirchen, Germany. Site‐ and species‐specific onset dates for the phenological phases were determined and related to elevation, temperature lapse rates and site‐specific temperature sums. Increasing temperatures induced advanced spring and delayed autumn phases, in which both yielded similar magnitudes. Delayed leaf senescence could therefore have been underestimated until now in extending the vegetation period. Not only the vegetation period, but also phenological periods extended with increasing temperature. Moreover, sensitivity to elevation and temperature strongly depends on the specific phenological phase. Differences between species and groups of species (deciduous, evergreen, high elevation) were found in onset dates, phenological response rates and also in the effect of chilling and forcing temperatures. Increased chilling days highly reduced forcing temperature requirements for deciduous trees, but less for evergreen trees. The problem of shifted species associations and phenological mismatches due to species‐specific responses to increasing temperature is a recent topic in ecological research. Therefore, we consider our findings from this novel, dense observation network in an alpine area of particular importance to deepen knowledge on phenological responses to climate change.     

Does flower phenology mirror the slowdown of global warming?

Although recent global warming trends in air temperature are not as pronounced as those observed only one decade ago, global mean temperature is still at a very high level. Does plant phenology – which is believed to be a suitable indicator of climate change – respond in a similar way, that is, does it still mirror recent temperature variations? We explored in detail long-term flowering onset dates of snowdrop, cherry, and lime tree and relevant spring temperatures at three sites in Germany (1901–2012) using the Bayesian multiple change-point approach. We investigated whether mean spring temperature changes were amplified or slowed down in the past decade and how plant phenology responded to the most recent temperature changes. Incorporating records with different end points (i.e., 2002 and 2012), we compared differences in trends and inferred possible differences caused by extrapolating phenological and meteorological data. The new multiple-change point approach is characterized by an enhanced structure and greater flexibility compared to the one change point model. However, the highest model probabilities for phenological (meteorological) records were still obtained for the one change point (linear) model. Marked warming trends in the recent decade were only revealed for mean temperatures of March to May, here better described with one or two change point models. In the majority of cases analyzed, changes in temperatures were well mirrored by phenological changes. However, temperatures in March to May were linked to less strongly advancing onset dates for lime tree flowering during the period 1901-2012, pointing to the likely influence of photoperiodic constraints or unfulfilled chilling requirements. Due to the slowdown of temperature increase, analyses conducted on records ending in 2002 demonstrated distinct differences when compared with records ending in 2012. Extrapolation of trends could therefore (along with the choice of the statistical method) lead to distinctly different results and most recent data should be integrated in order not to over- or underestimate future phenological changes.