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.     

Partly decoupled tree-ring width and leaf phenology response to 20th century temperature change in Sweden

The recent warming trend, and associated shifts in growing season length, challenge the principle of uniformitarianism, i.e., that current relations are persistent over time, and complicates the uncritical inferences of past climate from tree-ring data. Here we conduct a comparison between tree-ring width chronologies of Pinus sylvestris L. (Scots pine), Picea abies (L.) Karst. (Norway spruce) and Betula pubescens Ehrh. (Downy birch) and phenological observations (budburst and leaf senescence) of Fagus sylvatica L. (European beech), Quercus robur L. (European oak), Betula sp. (Birch), Norway spruce and Scots pine) in Sweden to assess to what extent the tree-ring width–temperature relationship and the timing of phenological phases are affected by increased temperature. Daily meteorological observations confirm a prolongation of the thermal growing season, most consistently observed as an earlier onset of around 1–2 weeks since the beginning of the 20th century. Observations of budburst closely mimic this pattern, with budburst of the deciduous trees occurring 1–2.5 weeks earlier. In contrast to the changes seen in phenology and observational temperature data, the tree-ring width–temperature relationships remain surprisingly stable throughout the 20th century. Norway spruce, Scots pine and Downy birch all show consistently significant correlations with at least one 30 day-long window of temperature starting in late June–early July season. Norway spruce displays the largest degree of stability, with a consistent 60 day-long temperature window with significant correlation starting around Julian calendar day 150. Thus, our results suggest that the principle of uniformitarianism is not violated during the period covered by modern meteorological observations. Further research is needed to determine at what thresholds the temperature sensitivity of these species may alter or deteriorate as a consequence of the ongoing climate change.     

Trends and uncertainties in budburst projections of Norway spruce in Northern Europe

Budburst is regulated by temperature conditions, and a warming climate is associated with earlier budburst. A range of phenology models has been developed to assess climate change effects, and they tend to produce different results. This is mainly caused by different model representations of tree physiology processes, selection of observational data for model parameterization, and selection of climate model data to generate future projections. In this study, we applied (i) Bayesian inference to estimate model parameter values to address uncertainties associated with selection of observational data, (ii) selection of climate model data representative of a larger dataset, and (iii) ensembles modeling over multiple initial conditions, model classes, model parameterizations, and boundary conditions to generate future projections and uncertainty estimates. The ensemble projection indicated that the budburst of Norway spruce in northern Europe will on average take place 10.2 ± 3.7 days earlier in 2051–2080 than in 1971–2000, given climate conditions corresponding to RCP 8.5. Three provenances were assessed separately (one early and two late), and the projections indicated that the relationship among provenance will remain also in a warmer climate. Structurally complex models were more likely to fail predicting budburst for some combinations of site and year than simple models. However, they contributed to the overall picture of current understanding of climate impacts on tree phenology by capturing additional aspects of temperature response, for example, chilling. Model parameterizations based on single sites were more likely to result in model failure than parameterizations based on multiple sites, highlighting that the model parameterization is sensitive to initial conditions and may not perform well under other climate conditions, whether the change is due to a shift in space or over time. By addressing a range of uncertainties, this study showed that ensemble modeling provides a more robust impact assessment than would a single phenology model run.     

The timing of autumn senescence is affected by the timing of spring phenology: implications for predictive models

Autumn senescence regulates multiple aspects of ecosystem function, along with associated feedbacks to the climate system. Despite its importance, current understanding of the drivers of senescence is limited, leading to a large spread in predictions of how the timing of senescence, and thus the length of the growing season, will change under future climate conditions. The most commonly held paradigm is that temperature and photoperiod are the primary controls, which suggests a future extension of the autumnal growing season as global temperatures rise. Here, using two decades of ground‐ and satellite‐based observations of temperate deciduous forest phenology, we show that the timing of autumn senescence is correlated with the timing of spring budburst across the entire eastern United States. On a year‐to‐year basis, an earlier/later spring was associated with an earlier/later autumn senescence, both for individual species and at a regional scale. We use the observed relationship to develop a novel model of autumn phenology. In contrast to current phenology models, this model predicts that the potential response of autumn phenology to future climate change is strongly limited by the impact of climate change on spring phenology. Current models of autumn phenology therefore may overpredict future increases in the length of the growing season, with subsequent impacts for modeling future CO₂ uptake and evapotranspiration.     

Spatial heterogeneity in the timing of birch budburst in response to future climate warming in Ireland

In order to predict the impact of future climate warming on trees it is important to quantify the effect climate has on their development. Our understanding of the phenological response to environmental drivers has given rise to various mathematical models of the annual growth cycle of plants. These models simulate the timing of phenophases by quantifying the relationship between development and its triggers, typically temperature. In addition, other environmental variables have an important role in determining the timing of budburst. For example, photoperiod has been shown to have a strong influence on phenological events of a number of tree species, including Betula pubescens (birch). A recently developed model for birch (DORMPHOT), which integrates the effects of temperature and photoperiod on budburst, was applied to future temperature projections from a 19-member ensemble of regional climate simulations (on a 25 km grid) generated as part of the ENSEMBLES project, to simulate the timing of birch budburst in Ireland each year up to the end of the present century. Gridded temperature time series data from the climate simulations were used as input to the DORMPHOT model to simulate future budburst timing. The results showed an advancing trend in the timing of birch budburst over most regions in Ireland up to 2100. Interestingly, this trend appeared greater in the northeast of the country than in the southwest, where budburst is currently relatively early. These results could have implications for future forest planning, species distribution modeling, and the birch allergy season.     

Nucleotide polymorphisms related to altitude and physiological traits in contrasting provenances of Norway spruce (<Emphasis Type="Italic">Picea abies</Emphasis>)

Variation of sequences of six EST-derived markers was investigated in three Norway spruce (Picea abies [L.] Karst.) provenances originating from different altitudes growing at two contrasting trial plots in Slovakia (Veľky Lom 450 m a.s.l., Mútne-Zákamenné 1,250 m a.s.l.) within a spin-off experiment of the IUFRO 1964/68 Inventory Provenance Experiment with Norway spruce. Single nucleotide polymorphisms (SNP) were identified and differences in allele frequencies at polymorphic sites were tested against altitude or associated with physiological and growth traits (chlorophyll a fluorescence, frost resistance, height, diameter, budburst phenology).     

Bayesian calibration of the Unified budburst model in six temperate tree species

Numerous phenology models developed to predict the budburst date of trees have been merged into one Unified model (Chuine, 2000, J. Theor. Biol. 207, 337–347). In this study, we tested a simplified version of the Unified model (Unichill model) on six woody species. Budburst and temperature data were available for five sites across Belgium from 1957 to 1995. We calibrated the Unichill model using a Bayesian calibration procedure, which reduced the uncertainty of the parameter coefficients and quantified the prediction uncertainty. The model performance differed among species. For two species (chestnut and black locust), the model showed good performance when tested against independent data not used for calibration. For the four other species (beech, oak, birch, ash), the model performed poorly. Model performance improved substantially for most species when using site-specific parameter coefficients instead of across-site parameter coefficients. This suggested that budburst is influenced by local environment and/or genetic differences among populations. Chestnut, black locust and birch were found to be temperature-driven species, and we therefore analyzed the sensitivity of budburst date to forcing temperature in those three species. Model results showed that budburst advanced with increasing temperature for 1–3 days °C⁻¹, which agreed with the observed trends. In synthesis, our results suggest that the Unichill model can be successfully applied to chestnut and black locust (with both across-site and site-specific calibration) and to birch (with site-specific calibration). For other species, temperature is not the only determinant of budburst and additional influencing factors will need to be included in the model.     

Warming counteracts defoliation‐induced mismatch by increasing herbivore‐plant phenological synchrony

Climate change is altering phenology; however, the magnitude of this change varies among taxa. Compared with phenological mismatch between plants and herbivores, synchronization due to climate has been less explored, despite its potential implications for trophic interactions. The earlier budburst induced by defoliation is a phenological strategy for plants against herbivores. Here, we tested whether warming can counteract defoliation‐induced mismatch by increasing herbivore‐plant phenological synchrony. We compared the larval phenology of spruce budworm and budburst in balsam fir, black spruce, and white spruce saplings subjected to defoliation in a controlled environment at temperatures of 12, 17, and 22°C. Budburst in defoliated saplings occurred 6–24 days earlier than in the controls, thus mismatching needle development from larval feeding. This mismatch decreased to only 3–7 days, however, when temperatures warmed by 5 and 10°C, leading to a resynchronization of the host with spruce budworm larvae. The increasing synchrony under warming counteracts the defoliation‐induced mismatch, disrupting trophic interactions and energy flow between forest ecosystem and insect populations. Our results suggest that the predicted warming may improve food quality and provide better growth conditions for larval development, thus promoting longer or more intense insect outbreaks in the future.     

Climatic response of budburst in the mountain birch at two areas in northern Fennoscandia and possible responses to global change

The relationship between the climate and budburst of the mountain birch was evaluated for two areas in subarctic (ca 69°N) Fennoscandia; at Abisko, Swedish Lapland, and at Kevo, Finnish Lapland. Thermal time (TT, degree‐day sums) to budburst was calculated for experimental conditions in the laboratory and for in situ observations of budburst. Two types of models predicting leaf emergence in situ were used: 1) TT to budburst for different threshold temperatures based on daily mean (TT_Mean) or daily maximum (TT_Max) temperatures and 2) ecophysiological budburst models. The obtained models were used to estimate effects of a changed climate.
Laboratory experiments of TT to budburst indicated no differences in the thermal requirements at the two areas. Temperature requirements of budburst declined successively during the progression of spring, from ca 250 degree‐days (>+2°C) in January to ca 100 in May. No significant trend in the date of budburst was found over the last 70 (Abisko) or 20 (Kevo) yr. There were some differences in the type of model that best explained the date of budburst in situ at the two areas. For Kevo the best prediction (minimum root mean square error, RMSE) of budburst was obtained by a simple thermal time model (TT_Mean>5.5°C) from 1 January (RMSE=2.1). For budburst at Abisko, models based on daily maximum temperature fitted better than those based on daily means. For Abisko, models based on thermal time accumulation only showed systematic errors in the predicted budburst that were correlated with budburst previous year (BB_PY). Including this apparent memory effect in the model decreased the error by 2.4 d. The best prediction for Abisko was thus obtained using TT_MAX>6.5 (RMSE=3.1) from 1 January.
Using these models to predict the effect of a changed air temperature climate indicate 3–8 d earlier budburst for a one‐degree increase in temperature, the effect being smaller for Kevo than for Abisko. For both areas a change in May temperature has a larger effect on the date of budburst than changes in any other month.     

Drought response of five conifer species under contrasting water availability suggests high vulnerability of Norway spruce and European larch

The ability of tree species to cope with anticipated decrease in water availability is still poorly understood. We evaluated the potential of Norway spruce, Scots pine, European larch, black pine, and Douglas‐fir to withstand drought in a drier future climate by analyzing their past growth and physiological responses at a xeric and a mesic site in Central Europe using dendroecological methods. Earlywood, latewood, and total ring width, as well as the δ¹³C and δ¹⁸O in early‐ and latewood were measured and statistically related to a multiscalar soil water deficit index from 1961 to 2009. At the xeric site, δ¹³C values of all species were strongly linked to water deficits that lasted longer than 11 months, indicating a long‐term cumulative effect on the carbon pool. Trees at the xeric site were particularly sensitive to soil water recharge in the preceding autumn and early spring. The native species European larch and Norway spruce, growing close to their dry distribution limit at the xeric site, were found to be the most vulnerable species to soil water deficits. At the mesic site, summer water availability was critical for all species, whereas water availability prior to the growing season was less important. Trees at the mesic were more vulnerable to water deficits of shorter duration than the xeric site. We conclude that if summers become drier, trees growing on mesic sites will undergo significant growth reductions, whereas at their dry distribution limit in the Alps, tree growth of the highly sensitive spruce and larch may collapse, likely inducing dieback and compromising the provision of ecosystem services. However, the magnitude of these changes will be mediated strongly by soil water recharge in winter and thus water availability at the beginning of the growing season.     

Factors that determine the severity of <Emphasis Type="Italic">Betula</Emphasis> spp. pollen seasons in Poland (Poznań and Krakow) and the United Kingdom (Worcester and London)

The aim of this paper is to analyse variations in the severity of Betula pollen seasons, particularly in relation to meteorological parameters at four sites, Poznań and Krakow in Poland, and Worcester and London in the United Kingdom. Results show that there is a significant relationship between Betula pollen season severity and weather conditions both in the year before pollination and in the same year that pollen is released from the plant. Furthermore, it is likely that the magnitude of birch pollen seasons in Poznań, Worcester and London is linked in some way to different phases of the North Atlantic Oscillation (NAO). Significant positive relationships exist between birch pollen counts at Poznań and temperatures, rainfall and averages of the NAO in the year before pollination. An opposite relationship is evident at the two sites studied in the United Kingdom. There were significant positive correlations between the severity of birch pollen seasons recorded at Worcester and temperatures and averages of the NAO during the winter and spring in the year of pollination, and negative correlations at both Worcester and London with similar variables from the previous year. In addition, Betula pollen seasons in Krakow do not appear to be influenced by the NAO, which is probably the result of Krakow having a more continental climate.     

Genotypic variation in Norway spruce correlates to fungal communities in vegetative buds

The taxonomically diverse phyllosphere fungi inhabit leaves of plants. Thus, apart from the fungi's dispersal capacities and environmental factors, the assembly of the phyllosphere community associated with a given host plant depends on factors encoded by the host's genome. The host genetic factors and their influence on the assembly of phyllosphere communities under natural conditions are poorly understood, especially in trees. Recent work indicates that Norway spruce (Picea abies) vegetative buds harbour active fungal communities, but these are hitherto largely uncharacterized. This study combines internal transcribed spacer sequencing of the fungal communities associated with dormant vegetative buds with a genome‐wide association study (GWAS) in 478 unrelated Norway spruce trees. The aim was to detect host loci associated with variation in the fungal communities across the population, and to identify loci correlating with the presence of specific, latent, pathogens. The fungal communities were dominated by known Norway spruce phyllosphere endophytes and pathogens. We identified six quantitative trait loci (QTLs) associated with the relative abundance of the dominating taxa (i.e., top 1% most abundant taxa). Three additional QTLs associated with colonization by the spruce needle cast pathogen Lirula macrospora or the cherry spruce rust (Thekopsora areolata) in asymptomatic tissues were detected. The identification of the nine QTLs shows that the genetic variation in Norway spruce influences the fungal community in dormant buds and that mechanisms underlying the assembly of the communities and the colonization of latent pathogens in trees may be uncovered by combining molecular identification of fungi with GWAS.     

Predicting spatial and temporal patterns of bud‐burst and spring frost risk in north‐west Europe: the implications of local adaptation to climate

The timing of spring bud‐burst and leaf development in temperate, boreal and Arctic trees and shrubs fluctuates from year to year, depending on meteorological conditions. Over several generations, the sensitivity of bud‐burst to meteorological conditions is subject to selection pressure. The timing of spring bud‐burst is considered to be under opposing evolutionary pressures; earlier bud‐burst increases the available growing season (capacity adaptation) but later bud‐burst decreases the risk of frost damage to actively growing parts (survival adaptation). The optimum trade‐off between these two forms of adaptation may be considered an evolutionarily stable strategy that maximizes the long‐term ecological fitness of a phenotype under a given climate. Rapid changes in climate, as predicted for this century, are likely to exceed the rate at which trees and shrubs can adapt through evolution or migration. Therefore the response of spring phenology will depend not only on future climatic conditions but also on the limits imposed by adaptation to current and historical climate. Using a dataset of bud‐burst dates from twenty‐nine sites in Finland for downy birch (Betula pubescens Ehrh.), we parameterize a simple thermal time bud‐burst model in which the critical temperature threshold for bud‐burst is a function of recent historical climatic conditions and reflects a trade‐off between capacity and survival adaptation. We validate this approach with independent data from eight independent sites outside Finland, and use the parameterized model to predict the response of bud‐burst to future climate scenarios in north‐west Europe. Current strategies for budburst are predicted to be suboptimal for future climates, with bud‐burst generally occurring earlier than the optimal strategy. Nevertheless, exposure to frost risk is predicted to decrease slightly and the growing season is predicted to increase considerably across most of the region. However, in high‐altitude maritime regions exposure to frost risk following bud‐burst is predicted to increase.     

Silver fir and Douglas fir are more tolerant to extreme droughts than Norway spruce in south‐western Germany

Improving our understanding of the potential of forest adaptation is an urgent task in the light of predicted climate change. Long‐term alternatives for susceptible yet economically important tree species such as Norway spruce (Picea abies) are required, if the frequency and intensity of summer droughts will continue to increase. Although Silver fir (Abies alba) and Douglas fir (Pseudotsuga menziesii) have both been described as drought‐tolerant species, our understanding of their growth responses to drought extremes is still limited. Here, we use a dendroecological approach to assess the resistance, resilience, and recovery of these important central Europe to conifer species the exceptional droughts in 1976 and 2003. A total of 270 trees per species were sampled in 18 managed mixed‐species stands along an altitudinal gradient (400–1200 m a.s.l.) at the western slopes of the southern and central Black Forest in southwest Germany. While radial growth in all species responded similarly to the 1976 drought, Norway spruce was least resistant and resilient to the 2003 summer drought. Silver fir showed the overall highest resistance to drought, similarly to Douglas fir, which exhibited the widest growth rings. Silver fir trees from lower elevations were more drought prone than trees at higher elevations. Douglas fir and Norway spruce, however, revealed lower drought resilience at higher altitudes. Although the 1976 and 2003 drought extremes were quite different, Douglas fir maintained consistently the highest radial growth. Although our study did not examine population‐level responses, it clearly indicates that Silver fir and Douglas fir are generally more resistant and resilient to previous drought extremes and are therefore suitable alternatives to Norway spruce; Silver fir more so at higher altitudes. Cultivating these species instead of Norway spruce will contribute to maintaining a high level of productivity across many Central European mountain forests under future climate change.     

Impact of global warming on the tree species composition of boreal forests in Finland and effects on emissions of isoprenoids

This study aims to identify how climate change may influence total emissions of monoterpene and isoprene from boreal forest canopies. The whole of Finland is assumed to experience an annual mean temperature (T) increase of 4 °C and a precipitation increase of 10% by the year 2100. This will increase forest resources throughout the country. At the same time, the proportions of Scots pine (Pinus sylvestris) and Norway spruce (Picea abies) in southern Finland (60°≤ latitude < 65°N) will be reduced from the current 40–50% to less than 10–20%, with increased dominance of birches (Betula pendula and Betula pubescens). In northern Finland (65°≤ latitude < 70°N), the proportions of Norway spruce and Scots pine will be balanced at a level of about 40% as the result of an increase in Norway spruce from the current 21% to 37% and a concurrent reduction in Scots pine from 63% to 40%. The proportion of birches is predicted to increase from the current 17% to 23%, but these will become the dominant species only on the most fertile sites. Total mean emissions of monoterpene by Scots pine will be reduced by 80% in southern Finland, but will increase by 62% in the north. Emissions from Norway spruce canopies will increase by 4% in the south but by 428% in the north, while those from birch canopies will increase by about 300% and 113%, respectively. Overall emissions of monoterpene over the whole country amount to about 950 kg km^?2 y^?1 under current temperature conditions and will increase by 17% to 1100 kg km^?2 y^?1 with elevated temperature and precipitation, mainly because of an increase at northern latitudes. Under current conditions, emissions of isoprene follow the spatial distribution of spruce canopies (the only isoprene‐emitting tree species that forms forests in Finland) with four times higher emissions in the south than in the north. The elevated temperature and the changes in the areal distribution of Norway spruce will result in increases in isoprene emissions of about 37% in southern Finland and 435% in northern Finland. Annual mean isoprene emissions from Norway spruce canopies over the whole country will increase by about 60% up to the year 2100.     

On the state of the public ill health: premature mortality in the United Kingdom and Europe.

Recently published data on mortality in the European Economic Community and Scandinavia convincingly showed that mortality among men and women aged 45-64 was considerably higher in the United Kingdom than elsewhere. This applied to deaths due to circulatory and respiratory disease, cancer, and all causes. For example, in 1980 in Scotland twice as many, or more, women aged 55-64 per 100 000 died of heart disease than in Belgium, Denmark, France, Greece, West Germany, the Netherlands, Norway, and Sweden. Reductions in mortality from all causes during 1950-80 in the United Kingdom did not match those in other countries, such as Finland and France. Whether the public in the United Kingdom knows about its relatively poor mortality state is doubtful. To secure improved funding of appropriate preventive and treatment services directed at reducing premature mortality, public awareness should be raised urgently so that politicians and political parties will respond quickly in a way that the problem demands.     

GLOBE students, teachers, and scientists demonstrate variable differences between urban and rural leaf phenology

The urban heat island effect, classically associated with high impervious surface area (ISA), low vegetation fractional cover (Fr), and high land surface temperature (LST), has been linked to changing patterns of vegetation phenology, especially spring growth. In this study, a collaboration with the Global Learning and Observations to Benefit the Environment (GLOBE) program, we investigated the effect of the urban environment on the timing of leaf budburst of native deciduous trees in seven cities: Asia (Tokyo, Japan; Bangkok and Korat, Thailand), Europe (Jyväskylä, Finland; Bishkek, Kyrgyzstan), Africa (Dakar, Senegal), and North America (Fairbanks, Alaska). The cities differed not only in population size but also in climate and vegetation type. Using Landsat satellite imagery from each city, we calculated LST, Fr, and ISA, and classified sites within each study area as rural or urban. The timing of leaf flushing, measured by students using GLOBE budburst protocols, was statistically different within all cities, with absolute differences ranging from 1 to 23 days. We assessed the classic urban phenology paradigm, which proposes higher LST, lower Fr, and earlier budburst in urban areas of temperate cities. Of the four temperate cities, Tokyo followed the classic paradigm, but no other city demonstrated consistent support. Urban budburst was advanced in three of the four temperate cities, but in only one of the three tropical cities. Results suggest that while vegetation phenology is consistently different between urban and rural areas, a uniform paradigm based on the explanatory variables in this study did not emerge. Although not testable here, it is likely that alterations to chilling requirements in temperate climates and humidity in tropical climates may also influence observed budburst differences.     

Synchronisation of egg hatching of brown hairstreak (Thecla betulae) and budburst of blackthorn (Prunus spinosa) in a warmer future

Synchronisation of the phenology of insect herbivores and their larval food plant is essential for the herbivores’ fitness. The monophagous brown hairstreak (Thecla betulae) lays its eggs during summer, hibernates as an egg, and hatches in April or May in the Netherlands. Its main larval food plant blackthorn (Prunus spinosa) flowers in early spring, just before the leaves appear. As soon as the Blackthorn opens its buds, and this varies with spring temperatures, food becomes available for the brown hairstreak. However, the suitability of the leaves as food for the young caterpillars is expected to decrease rapidly. Therefore, the timing of egg hatch is an important factor for larval growth. This study evaluates food availability for brown hairstreak at different temperatures. Egg hatch and budburst were monitored from 2004 to 2008 at different sites in the Netherlands. Results showed ample food availability at all monitored temperatures and sites but the degree of synchrony varied strongly with spring temperatures. To further study the effect of temperature on synchronisation, an experiment using normal temperatures of a reference year (T) and temperatures of T + 5°C was carried out in climate chambers. At T + 5°C, both budburst and egg hatch took place about 20 days earlier and thus, on average, elevated temperature did not affect synchrony. However, the total period of budburst was 11 days longer, whereas the period of egg hatching was 3 days shorter. The implications for larval growth by the brown hairstreak under a warmer climate are considered.     

Leafing phenology and timing of egg laying in great tits Parus major and blue tits P. caeruleus

To evaluate the importance of tree leafing for the start of laying and clutch size of birds, we compared the breeding phenology of great tits Parus major and blue tits P. caeruleus between one coastal and two inland sites in the same geographical region. Because of the cooling influence of the sea, trees at the coastal site were known to initiate budburst about a week later than at the inland sites. During 5 years, breeding by the tits and the leaf phenology of birch Betula pendula , and oak Quercus robur were monitored. The leaf phenology of birch and oak explained a significant part of the between-year variation in the start of egg laying in blue and great tits, respectively. The tits started laying earlier at the sites with an early budburst, i.e. normally inland. However, leaf phenology was not an absolute cue to the start of laying, since blue tits laid earlier relative to leafing at the inland site than at the coastal site, and both tit species laid eggs earlier relative to leafing during late springs. In neither species was clutch size affected by leafing phenology. However, great tit females at the coastal site consistently produced fewer eggs than did those at the inland site. No such difference was found in the blue tits. Although leafing phenology may predict the start of laying in tits, other factors also influence its timing. These factors might include other cues, or differing life-history trade-offs depending on site or general climatic factors during the spring.     

Multiscale modeling of spring phenology across Deciduous Forests in the Eastern United States

Phenological events, such as bud burst, are strongly linked to ecosystem processes in temperate deciduous forests. However, the exact nature and magnitude of how seasonal and interannual variation in air temperatures influence phenology is poorly understood, and model‐based phenology representations fail to capture local‐ to regional‐scale variability arising from differences in species composition. In this paper, we use a combination of surface meteorological data, species composition maps, remote sensing, and ground‐based observations to estimate models that better represent how community‐level species composition affects the phenological response of deciduous broadleaf forests to climate forcing at spatial scales that are typically used in ecosystem models. Using time series of canopy greenness from repeat digital photography, citizen science data from the USA National Phenology Network, and satellite remote sensing‐based observations of phenology, we estimated and tested models that predict the timing of spring leaf emergence across five different deciduous broadleaf forest types in the eastern United States. Specifically, we evaluated two different approaches: (i) using species‐specific models in combination with species composition information to ‘upscale’ model predictions and (ii) using repeat digital photography of forest canopies that observe and integrate the phenological behavior of multiple representative species at each camera site to calibrate a single model for all deciduous broadleaf forests. Our results demonstrate variability in cumulative forcing requirements and photoperiod cues across species and forest types, and show how community composition influences phenological dynamics over large areas. At the same time, the response of different species to spatial and interannual variation in weather is, under the current climate regime, sufficiently similar that the generic deciduous forest model based on repeat digital photography performed comparably to the upscaled species‐specific models. More generally, results from this analysis demonstrate how in situ observation networks and remote sensing data can be used to synergistically calibrate and assess regional parameterizations of phenology in models.