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.     

A critical thermal transition driving spring phenology of Northern Hemisphere conifers

Despite growing interest in predicting plant phenological shifts, advanced spring phenology by global climate change remains debated. Evidence documenting either small or large advancement of spring phenology to rising temperature over the spatio-temporal scales implies a potential existence of a thermal threshold in the responses of forests to global warming. We collected a unique data set of xylem cell-wall-thickening onset dates in 20 coniferous species covering a broad mean annual temperature (MAT) gradient (−3.05 to 22.9°C) across the Northern Hemisphere (latitudes 23°–66° N). Along the MAT gradient, we identified a threshold temperature (using segmented regression) of 4.9 ± 1.1°C, above which the response of xylem phenology to rising temperatures significantly decline. This threshold separates the Northern Hemisphere conifers into cold and warm thermal niches, with MAT and spring forcing being the primary drivers for the onset dates (estimated by linear and Bayesian mixed-effect models), respectively. The identified thermal threshold should be integrated into the Earth-System-Models for a better understanding of spring phenology in response to global warming and an improved prediction of global climate-carbon feedbacks.     

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.     

Pattern of xylem phenology in conifers of cold ecosystems at the Northern Hemisphere

The interaction between xylem phenology and climate assesses forest growth and productivity and carbon storage across biomes under changing environmental conditions. We tested the hypothesis that patterns of wood formation are maintained unaltered despite the temperature changes across cold ecosystems. Wood microcores were collected weekly or biweekly throughout the growing season for periods varying between 1 and 13 years during 1998–2014 and cut in transverse sections for assessing the onset and ending of the phases of xylem differentiation. The data set represented 1321 trees belonging to 10 conifer species from 39 sites in the Northern Hemisphere and covering an interval of mean annual temperature exceeding 14 K. The phenological events and mean annual temperature of the sites were related linearly, with spring and autumnal events being separated by constant intervals across the range of temperature analysed. At increasing temperature, first enlarging, wall‐thickening and mature tracheids appeared earlier, and last enlarging and wall‐thickening tracheids occurred later. Overall, the period of wood formation lengthened linearly with the mean annual temperature, from 83.7 days at ?2 °C to 178.1 days at 12 °C, at a rate of 6.5 days °C^?1. April–May temperatures produced the best models predicting the dates of wood formation. Our findings demonstrated the uniformity of the process of wood formation and the importance of the environmental conditions occurring at the time of growth resumption. Under warming scenarios, the period of wood formation might lengthen synchronously in the cold biomes of the Northern Hemisphere.     

Phenology model from surface meteorology does not capture satellite-based greenup estimations

Seasonal temperature change in temperate forests is known to trigger the start of spring growth, and both interannual and spatial variations in spring onset have been tied to climatic variability. Satellite dates are increasingly being used in phenology studies, but to date that has been little effort to link remotely sensed phenology to surface climate records. In this research, we use a two‐parameter spring warming phenology model to explore the relationship between climate and satellite‐based phenology. We employ daily air temperature records between 2000 and 2005 for 171 National Oceanographic and Atmospheric Administration weather stations located throughout New England to construct spring warming models predicting the onset of spring, as defined by the date of half‐maximum greenness (D₅₀) in deciduous forests as detected from Moderate Resolution Imaging Spectrometer. The best spring warming model starts accumulating temperatures after March 20th and when average daily temperatures exceed 5°C. The accumulated heat sums [heating degree day (HDD)] required to reach D₅₀ range from 150 to 300 degree days over New England, with the highest requirements to the south and in coastal regions. We test the ability of the spring warming model to predict phenology against a null photoperiod model (average date of onset). The spring warming model offers little improvement on the null model when predicting D₅₀. Differences between the efficacies of the two models are expressed as the ‘climate sensitivity ratio’ (CSR), which displays coherent spatial patterns. Our results suggest that northern (beech‐maple‐birch) and central (oak‐hickory) hardwood forests respond to climate differently, particularly with disparate requirements for the minimum temperature necessary to begin spring growth (3 and 6°C, respectively). We conclude that spatial location and species composition are critical factors for predicting the phenological response to climate change: satellite observations cannot be linked directly to temperature variability if species or community compositions are unknown.     

Will changes in phenology track climate change? A study of growth initiation timing in coast Douglas‐fir

Under climate change, the reduction of frost risk, onset of warm temperatures and depletion of soil moisture are all likely to occur earlier in the year in many temperate regions. The resilience of tree species will depend on their ability to track these changes in climate with shifts in phenology that lead to earlier growth initiation in the spring. Exposure to warm temperatures (‘forcing’) typically triggers growth initiation, but many trees also require exposure to cool temperatures (‘chilling’) while dormant to readily initiate growth in the spring. If warming increases forcing and decreases chilling, climate change could maintain, advance or delay growth initiation phenology relative to the onset of favorable conditions. We modeled the timing of height‐ and diameter‐growth initiation in coast Douglas‐fir (an ecologically and economically vital tree in western North America) to determine whether changes in phenology are likely to track changes in climate using data from field‐based and controlled‐environment studies, which included conditions warmer than those currently experienced in the tree's range. For high latitude and elevation portions of the tree's range, our models predicted that warming will lead to earlier growth initiation and allow trees to track changes in the onset of the warm but still moist conditions that favor growth, generally without substantially greater exposure to frost. In contrast, toward lower latitude and elevation range limits, the models predicted that warming will lead to delayed growth initiation relative to changes in climate due to reduced chilling, with trees failing to capture favorable conditions in the earlier parts of the spring. This maladaptive response to climate change was more prevalent for diameter‐growth initiation than height‐growth initiation. The decoupling of growth initiation with the onset of favorable climatic conditions could reduce the resilience of coast Douglas‐fir to climate change at the warm edges of its distribution.     

Spring phenology at different altitudes is becoming more uniform under global warming in Europe

Under current global warming, high‐elevation regions are expected to experience faster warming than low‐elevation regions. However, due to the lack of studies based on long‐term large‐scale data, the relationship between tree spring phenology and the elevation‐dependent warming is unclear. Using 652k records of leaf unfolding of five temperate tree species monitored during 1951–2013 in situ in Europe, we discovered a nonlinear trend in the altitudinal sensitivity (S_A, shifted days per 100 m in altitude) in spring phenology. A delayed leaf unfolding (2.7 ± 0.6 days per decade) was observed at high elevations possibly due to decreased spring forcing between 1951 and 1980. The delayed leaf unfolding at high‐elevation regions was companied by a simultaneous advancing of leaf unfolding at low elevations. These divergent trends contributed to a significant increase in the S_A (0.36 ± 0.07 days 100/m per decade) during 1951–1980. Since 1980, the S_A started to decline with a rate of ?0.32 ± 0.07 days 100/m per decade, possibly due to reduced chilling at low elevations and improved efficiency of spring forcing in advancing the leaf unfolding at high elevations, the latter being caused by increased chilling. Our results suggest that due to both different temperature changes at the different altitudes, and the different tree responses to these changes, the tree phenology has shifted at different rates leading to a more uniform phenology at different altitudes during recent decades.     

Challenging a 15‐year‐old claim: The North Atlantic Oscillation index as a predictor of spring migration phenology of birds

Many migrant bird species that breed in the Northern Hemisphere show advancement in spring arrival dates. The North Atlantic Oscillation (NAO) index is one of the climatic variables that have been most often investigated and shown to be correlated with these changes in spring arrival. Although the NAO is often claimed to be a good predictor or even to have a marked effect on interannual changes in spring migration phenology of Northern Hemisphere breeding birds, the results on relations between spring migration phenology and NAO show a large variety, ranging from no, over weak, to a strong association. Several factors, such as geographic location, migration phase, and the NAO index time window, have been suggested to partly explain these observed differences in association. A combination of a literature meta‐analysis, and a meta‐analysis and sliding time window analysis of a dataset of 23 short‐ and long‐distance migrants from the constant‐effort trapping garden at Helgoland, Germany, however, paints a completely different picture. We found a statistically significant overall effect size of the NAO on spring migration phenology (coefficient = ?0.14, SE = 0.054), but this on average only explains 0%–6% of the variance in spring migration phenology across all species. As such, the value and biological meaning of the NAO as a general predictor or explanatory variable for climate change effects on migration phenology of birds, seems highly questionable. We found little to no definite support for previously suggested factors, such as geographic location, migration phenology phase, or the NAO time window, to explain the heterogeneity in correlation differences. We, however, did find compelling evidence that the lack of accounting for trends in both time series has led to strongly inflated (spurious) correlations in many studies (coefficient = ?0.13, SE = 0.019).     

Three‐dimensional change in temperature sensitivity of northern vegetation phenology

Understanding how the temperature sensitivity of phenology changes with three spatial dimensions (altitude, latitude, and longitude) is critical for the prediction of future phenological synchronization. Here we investigate the spatial pattern of temperature sensitivity of spring and autumn phenology with altitude, latitude, and longitude during 1982–2016 across mid‐ and high‐latitude Northern Hemisphere (north of 30°N). We find distinct spatial patterns of temperature sensitivity of spring phenology (hereafter “spring S_T”) among altitudinal, latitudinal, and longitudinal gradient. Spring S_T decreased with altitude mostly over eastern Europe, whereas the opposite occurs in eastern North America and the north China plain. Spring S_T decreased with latitude mainly in the boreal regions of North America, temperate Eurasia, and the arid/semi‐arid regions of Central Asia. This distribution may be related to the increased temperature variance, decreased precipitation, and radiation with latitude. Compared to spring S_T, the spatial pattern of temperature sensitivity of autumn phenology (hereafter “autumn S_T”) is more heterogeneous, only showing a clear spatial pattern of autumn S_T along the latitudinal gradient. Our results highlight the three‐dimensional view to understand the phenological response to climate change and provide new metrics for evaluating phenological models. Accordingly, establishing a dense, high‐quality three‐dimensional observation system of phenology data is necessary for enhancing our ability to both predict phenological changes under changing climatic conditions and to facilitate sustainable management of ecosystems.     

Regional unified model‐based leaf unfolding prediction from 1960 to 2009 across northern China

Using first leaf unfolding data of Salix matsudana, Populus simonii, Ulmus pumila, and Prunus armeniaca, and daily mean temperature data during the 1981–2005 period at 136 stations in northern China, we fitted unified forcing and chilling phenology models and selected optimum models for each species at each station. Then, we examined performances of each optimum local species‐specific model in predicting leaf unfolding dates at all external stations within the corresponding climate region and selected 16 local species‐specific models with maximum effective predictions as the regional unified models in different climate regions. Furthermore, we validated the regional unified models using leaf unfolding and daily mean temperature data beyond the time period of model fitting. Finally, we substituted gridded daily mean temperature data into the regional unified models, and reconstructed spatial patterns of leaf unfolding dates of the four tree species across northern China during 1960–2009. At local scales, the unified forcing model shows higher simulation efficiency at 83% of data sets, whereas the unified chilling model indicates higher simulation efficiency at 17% of data sets. Thus, winter temperature increase so far has not yet significantly influenced dormancy and consequent leaf development of deciduous trees in most parts of northern China. Spatial and temporal validation confirmed capability and reliability of regional unified species‐specific models in predicting leaf unfolding dates in northern China. Reconstructed leaf unfolding dates of the four tree species show significant advancements by 1.4–1.6 days per decade during 1960–2009 across northern China, which are stronger for the earlier than the later leaf unfolding species. Our findings suggest that the principal characteristics of plant phenology and phenological responses to climate change at regional scales can be captured by phenological and climatic data sets at a few representative locations.     

A new seasonal‐deciduous spring phenology submodel in the Community Land Model 4.5: impacts on carbon and water cycling under future climate scenarios

A spring phenology model that combines photoperiod with accumulated heating and chilling to predict spring leaf‐out dates is optimized using PhenoCam observations and coupled into the Community Land Model (CLM) 4.5. In head‐to‐head comparison (using satellite data from 2003 to 2013 for validation) for model grid cells over the Northern Hemisphere deciduous broadleaf forests (5.5 million km²), we found that the revised model substantially outperformed the standard CLM seasonal‐deciduous spring phenology submodel at both coarse (0.9 × 1.25°) and fine (1 km) scales. The revised model also does a better job of representing recent (decadal) phenological trends observed globally by MODIS, as well as long‐term trends (1950–2014) in the PEP725 European phenology dataset. Moreover, forward model runs suggested a stronger advancement (up to 11 days) of spring leaf‐out by the end of the 21st century for the revised model. Trends toward earlier advancement are predicted for deciduous forests across the whole Northern Hemisphere boreal and temperate deciduous forest region for the revised model, whereas the standard model predicts earlier leaf‐out in colder regions, but later leaf‐out in warmer regions, and no trend globally. The earlier spring leaf‐out predicted by the revised model resulted in enhanced gross primary production (up to 0.6 Pg C yr^?1) and evapotranspiration (up to 24 mm yr^?1) when results were integrated across the study region. These results suggest that the standard seasonal‐deciduous submodel in CLM should be reconsidered, otherwise substantial errors in predictions of key land–atmosphere interactions and feedbacks may result.     

Predicting xylem phenology in black spruce under climate warming

In the next century, the boreal ecosystems are projected to experience greater rates of warming than most other regions of the world. As the boreal forest constitutes a reservoir of trees of huge ecological importance and only partially known economic potential, any possible climate‐related change in plant growth and dynamics has to be promptly predicted and evaluated. A model for assessing xylem phenology in black spruce [Picea mariana (Mill.) B.S.P.] using daily temperatures and thermal thresholds was defined and applied to predict changes in onset, ending and duration of xylem growth under different warming scenarios with temperatures rising by up to 3 °C. This was achieved by collecting and analyzing a dataset obtained from a 7‐year monitoring of cambium phenology and wood formation on a weekly time‐scale in trees growing in four sites at different latitudes and altitudes in the Saguenay‐Lac‐Saint‐Jean region (Quebec, Canada). The onset of xylem growth occurred between mid‐May and early June while the end ranged between mid‐September and early October, resulting in a growing season of 101–141 days. The model predicted longer duration of xylem growth at higher temperatures, with an increase of 8–11 days/ °C, because of an earlier onset and later ending of growth. With an increase of 3 °C in the mean temperature during the year, the duration of xylem growth changed on average from 125 to 160 days. The predicted changes in cambial phenology could significantly affect future wood production of the boreal ecosystems.     

New insights on plant phenological response to temperature revealed from long‐term widespread observations in China

Constraints of temperature on spring plant phenology are closely related to plant growth, vegetation dynamics, and ecosystem carbon cycle. However, the effects of temperature on leaf onset, especially for winter chilling, are still not well understood. Using long‐term, widespread in situ phenology observations collected over China for multiple plant species, this study analyzes the quantitative response of leaf onset to temperature, and compares empirical findings with existing theories and modeling approaches, as implemented in 18 phenology algorithms. Results show that the growing degree days (GDD) required for leaf onset vary distinctly among plant species and geographical locations as well as at organizational levels (species and community), pointing to diverse adaptation strategies. Chilling durations (CHD) needed for releasing bud dormancy decline monotonously from cold to warm areas with very limited interspecies variations. Results also reveal that winter chilling is a crucial component of phenology models, and its effect is better captured with an index that accounts for the inhomogeneous effectiveness of low temperature to chilling rate than with the conventional CHD index. The impact of spring warming on leaf onset is nonlinear, better represented by a logistical function of temperature than by the linear function currently implemented in biosphere models. The optimized base temperatures for thermal accumulation and the optimal chilling temperatures are species‐dependent and average at 6.9 and 0.2°C, respectively. Overall, plants’ chilling requirement is not a constant, and more chilling generally results in less requirement of thermal accumulation for leaf onset. Our results clearly demonstrate multiple deficiencies of the parameters (e.g., base temperature) and algorithms (e.g., method for calculating GDD) in conventional phenology models to represent leaf onset. Therefore, this study not only advances our mechanistic and quantitative understanding of temperature controls on leaf onset but also provides critical information for improving existing phenology 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.     

Moisture‐driven xylogenesis in Pinus ponderosa from a Mojave Desert mountain reveals high phenological plasticity

Future seasonal dynamics of wood formation in hyperarid environments are still unclear. Although temperature‐driven extension of the growing season and increased forest productivity are expected for boreal and temperate biomes under global warming, a similar trend remains questionable in water‐limited regions. We monitored cambial activity in a montane stand of ponderosa pine (Pinus ponderosa) from the Mojave Desert for 2 consecutive years (2015–2016) showing opposite‐sign anomalies between warm‐ and cold‐season precipitation. After the wet winter/spring of 2016, xylogenesis started 2 months earlier compared to 2015, characterized by abundant monsoonal (July–August) rainfall and hyperarid spring. Tree size did not influence the onset and ending of wood formation, highlighting a predominant climatic control over xylem phenological processes. Moisture conditions in the previous month, in particular soil water content and dew point, were the main drivers of cambial phenology. Latewood formation started roughly at the same time in both years; however, monsoonal precipitation triggered the formation of more false rings and density fluctuations in 2015. Because of uncertainties in future precipitation patterns simulated by global change models for the Southwestern United States, the dependency of P. ponderosa on seasonal moisture implies a greater conservation challenge than for species that respond mostly to temperature conditions.     

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.     

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.     

Hydraulic acclimation to shading in boreal conifers of varying shade tolerance

The purpose of this study was to determine how shading affects the hydraulic and wood‐anatomical characteristics of four boreal conifers (Pinus banksiana, Pinus contorta, Picea glauca and Picea mariana) that differ in shade tolerance. Plants were grown in an open field and under a deciduous‐dominated overstory for 6 years. Sapwood‐ and leaf‐area specific conductivity, vulnerability curves, and anatomical measurements (light and scanning electron microscopy) were made on leading shoots from six to nine trees of each treatment combination. There was no difference in sapwood‐area specific conductivity between open‐grown and understory conifers, although two of four species had larger tracheid diameters in the open. Shaded conifers appeared to compensate for small diameter tracheids by changes in pit membrane structure. Scanning electron microscopy revealed that understory conifers had thinner margo strands, greater maximum pore size in the margo, and more torus extensions. All of these trends may contribute to inadequate sealing of the torus. This is supported by the fact that all species showed increased vulnerability to cavitation when grown in the understory. Although evaporative demand in an understory environment is low, a rapid change into fully exposed conditions could be detrimental for shaded conifers.     

High‐resolution prediction of leaf onset date in Japan in the 21st century under the IPCC A1B scenario

Reports indicate that leaf onset (leaf flush) of deciduous trees in cool‐temperate ecosystems is occurring earlier in the spring in response to global warming. In this study, we created two types of phenology models, one driven only by warmth (spring warming [SW] model) and another driven by both warmth and winter chilling (parallel chill [PC] model), to predict such phenomena in the Japanese Islands at high spatial resolution (500 m). We calibrated these models using leaf onset dates derived from satellite data (Terra/MODIS) and in situ temperature data derived from a dense network of ground stations Automated Meteorological Data Acquisition System. We ran the model using future climate predictions created by the Japanese Meteorological Agency's MRI‐AGCM3.1S model. In comparison to the first decade of the 2000s, our results predict that the date of leaf onset in the 2030s will advance by an average of 12 days under the SW model and 7 days under the PC model throughout the study area. The date of onset in the 2090s will advance by 26 days under the SW model and by 15 days under the PC model. The greatest impact will occur on Hokkaido (the northernmost island) and in the central mountains.     

The effects of chilling and forcing temperatures on spring synchrony between larch casebearer and tamarack

1. Spring phenological synchrony can be important for tree-insect interactions. Depending on the magnitude and direction of phenological shifts, overwintering insects could be affected in many ways, for example, facing starvation or having to contend with increased chemical or physical defences of host trees. If temperature has different influences on the phenology of trees and insects, climate change can alter spring phenological synchrony.
2. In this experiment, we exposed tamarack seedlings and larch case bearer larvae from Minnesota, USA, to a variety of chilling and forcing temperatures and measured spring phenology (twig bud break and larval activation). We additionally measured case bearer performance on seedlings that were exposed to different forcing × chilling levels, tracking larval survivorship to adulthood.
3. Warmer forcing enhanced larval activation and bud break, but larval development slowed down past 21°C. Higher chilling temperatures accelerated bud break, but the effect was inconclusive for larvae. There was no chilling × forcing interaction for either species. Spring activity accelerated more quickly with increases in temperature for larvae than for seedlings, resulting in increased phenological synchrony at warmer temperatures. Activation rates for overwintering larvae were highest at 27°C, while survivorship to adulthood following spring activation was highest at 21°C. At temperatures at or beyond 27°C, no larvae reached adulthood.
4. Warmer winters and springs will likely initially increase spring synchrony between tamarack and larch case bearer, exposing larvae to younger, potentially more nutritious foliage, but extremely warm spring temperatures may decrease survivorship of larvae to adulthood.