Advancing Spring Phenology Research Articles

Duration and variability of spring green-up mediate population consequences of climate change.

Single phenological measures, like the average rate of phenological advancement, may be insufficient to explain how climate change is driving trends in animal populations. Here, we develop a multifactorial concept of spring phenology-including the onset of spring, spring duration, interannual variability, and their temporal changes-as a driver for population dynamics of migratory terrestrial species in seasonal environments. Using this conceptual model, we found that effects of advancing spring phenology on animal populations may be buffered or amplified depending on the duration and interannual variability of spring green-up, and those effects are modified by evolutionary and plastic adaptations of species. Furthermore, we compared our modelling results with empirical data on normalized difference vegetation index-based spring green-up phenology and population trends of 106 European landbird finding similar associations. We conclude how phenological changes are expected to affect migratory bird populations across Europe and identify regions that are particularly prone to suffer population declines.

Effects of Spring Phenology on Grassland Growth in Qilian Mountains Across Multiple Spatiotemporal Scales

AbstractInvestigating the differences in grassland growth and its response to advancing spring phenology and climate change across various spatiotemporal scales is crucial for the sustainable development of alpine mountain regions under a warming and wetting climate. This study analyzed the growth dynamics of grassland in the Qilian Mountains from 2000 to 2019 at seasonal and periodic scales. Using partial wavelet coherency and partial correlation analysis, we investigated the influence of temperature, precipitation, and spring phenology on grassland growth. Additionally, we quantified the dominant factors affecting grassland growth in different ecological zones and elevational gradient. The results indicated that the mean normalized difference vegetation index increased at an average rate of 1.1 and 2.8 × 10−3/year for grassland in spring and summer, respectively. Furthermore, the start of vegetation growing season (SOS) has advanced 6.0 days during 2000–2019. The dominant factor for grassland growth shifted from spring temperature to summer precipitation. The importance of SOS ranked second, significantly influencing 29.3% and 25.1% of grassland during spring and summer, respectively. Grasslands situated above 4,000 m above sea level (ASL) were more sensitive to climate change, with SOS exerting the greatest influence within the elevation range of 2,800–4,000 m ASL. Additionally, the phase difference in the time‐frequency scale of partial wavelet coherency revealed a lag effect of approximately 1 month for summer grassland growth to respond to precipitation. These findings can provide a scientific basis for optimizing global vegetation models and predicting the productivity of alpine mountain grasslands.

Urban environments provide new perspectives for forecasting vegetation phenology responses under climate warming.

Given that already-observed temperature increase within cities far exceeds the projected global temperature rise by the end of the century, urban environments often offer a unique opportunity for studying ecosystem response to future warming. However, the validity of thermal gradients in space serving as a substitute for those in time is rarely tested. Here, we investigated vegetation phenology dynamics in China's 343 cities and empirically test whether phenological responses to spatial temperature rise in urban settings can substitute for those to temporal temperature rise in their natural counterparts based on satellite-derived vegetation phenology and land surface temperature from 2003 to 2018. We found prevalent advancing spring phenology with "high confidence" and delaying autumn phenology with "medium confidence" under the context of widespread urban warming. Furthermore, we showed that space cannot substitute for time in predicting phenological shifts under climate warming at the national scale and for most cities. The thresholds of ~11°C mean annual temperature and ~600 mm annual precipitation differentiated the magnitude of phenological sensitivity to temperature across space and through time. Below those thresholds, there existed stronger advanced spring phenology and delayed autumn phenology across the spatial urbanization gradients than through time, and vice versa. Despite the complex and diverse relationships between phenological sensitivities across space and through time, we found that the directions of the temperature changes across spatial gradients were converged (i.e., mostly increased), but divergent through temporal gradients (i.e., increased or decreased without a predominant direction). Similarly, vegetation phenology changes more uniformly over space than through time. These results suggested that the urban environments provide a real-world condition to understand vegetation phenology response under future warming.

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.

The early bud gets the cold: Diverging spring phenology drives exposure to late frost in a Picea mariana [(Mill.) BSP] common garden.

Under climate change, the increasing occurrence of late frost combined with advancing spring phenology can increase the risk of frost damage in trees. In this study, we tested the link between intra-specific variability in bud phenology and frost exposure and damages. We analysed the effects of the 2021 late frost event in a black spruce (Picea mariana (Mill.) BSP) common garden in Québec, Canada. We hypothesised that the timing of budbreak drives the exposure of vulnerable tissues and explains differences in frost damage. Budbreak was monitored from 2015 to 2021 in 371 trees from five provenances originating between 48° and 53° N and planted in a common garden at 48° N. Frost damages were assessed on the same trees through the proportion of damaged buds per tree and related to the phenological phases by ordinal regressions. After an unusually warm spring, minimum temperatures fell to -1.9°C on May 28 and 29, 2021. At this moment, trees from the northern provenances were more advanced in their phenology and showed more frost damage. Provenances with earlier budbreak had a higher probability of damage occurrence according to ordinal regression. Our study highlights the importance of intra-specific variability of phenological traits on the risk of frost exposure. We provide evidence that the timings of bud phenology affect sensitivity to frost, leading to damages at temperatures of -1.9°C. Under the same conditions, the earlier growth reactivation observed in the northern provenances increases the risks of late frost damage on the developing buds.

Pilot Study of Sap Properties of Norway Spruce (Picea abies (L.) Karst.) Trees Used and Not Used for Sap-Feeding by Three-Toed Woodpeckers (Picoides tridactylus)

The ecophysiology of the Norway spruce (Picea abies (L.) Karst.) trees that were used by three-toed woodpeckers (TTW) (Picoides tridactylus) for their sap-feeding activities were investigated. The pilot study was conducted in southern Finland (61°15′ N, 25°00′ E). During April–June 2015, three different tree categories of Norway spruce were selected for monitoring: trees that were frequently used by the TTWs for phloem sap-feeding for several previous years; trees that were only recently used by TTWs for sap-feeding; and control trees that were not used at all for sap-feeding. Phloem sap and phloem tissue samples were frequently extracted from tree trunks and analyzed for the content and composition of nonstructural carbohydrates, phloem sap osmolality, solute, and water content, as well as for the content and composition of secondary metabolites typical for defense reactions in the phloem. Simple crown characteristics were also measured, including tree height, diameter at breast height, and their ratio (slenderness index). According to our results, the TTWs preferred Norway spruce trees that showed advanced spring phenology to feed on, as evidenced especially by the lower ratio of raffinose (typically high during the winter months) to total soluble sugars of phloem tissues as compared to non-used control trees. The lower slenderness index of the trees chosen by the TTWs indicates low canopy competition pressure with good access to light (i.e., the sun heats the trunks well in spring). There were no differences in the phloem osmolality or solute content between the used or unused control trees. The trees used by the TTWs had significantly higher concentrations of antioxidant phenolic (+)-catechins and stilbene glycosides in phloem tissue, and the stilbene content was also higher in the extracted sap. The phenolics content of the phloem tissue had a clear seasonal trend, being the highest in the early spring and lower towards the onset of the cambial growth processes. The phloem sap is rich in antioxidants and soluble sugars that are potentially beneficial for the TTWs, but more quantitative research is needed to explore the importance of the sap properties to TTWs.

Disorder or a new order: How climate change affects phenological variability.

Advancing spring phenology is a well documented consequence of anthropogenic climate change, but it is not well understood how climate change will affect the variability of phenology year to year. Species' phenological timings reflect the adaptation to a broad suite of abiotic needs (e.g., thermal energy) and biotic interactions (e.g., predation and pollination), and changes in patterns of variability may disrupt those adaptations and interactions. Here, we present a geographically and taxonomically broad analysis of phenological shifts, temperature sensitivity, and changes in interannual variability encompassing nearly 10,000 long-term phenology time series representing more than 1000 species across much of the Northern Hemisphere. We show that the timings of leaf-out, flowering, insect first-occurrence, and bird arrival were the most sensitive to temperature variation and have advanced at the fastest pace for early-season species in colder and less seasonal regions. We did not find evidence for changing variability in warmer years in any phenophase groups, although leaf-out and flower phenology have become moderately but significantly less variable over time. Our findings suggest that climate change has not to this point fundamentally altered the patterns of interannual phenological variability.

Impacts of climate change on vegetation phenology over the Great Lakes Region of Central Asia from 1982 to 2014

Dryland ecosystems in the Great Lakes Region of Central Asia (GLRCA) are highly sensitive to climate change due to the climate of spring precipitation. Although shifts in vegetation phenology have been widely attributed to rising temperature, the effects of solar radiation and drought on phenology remain largely unknown. Understanding the mechanisms of vegetation phenology response to climatic factors is essential for assessing the impact of climate change on dryland ecosystems. In this study, we investigated the spatial and temporal variations of vegetation phenology across the GLRCA using a long-term series of Normalized Difference Vegetation Index (NDVI), and then examined the response of vegetation phenology to climate change within different climate zones by combining with climate data (surface temperature, soil moisture, short-wave radiation, and standardized precipitation evapotranspiration index (SPEI)). The results suggested that the start of growing season (SGS) and the end of growing season (EGS) were significantly earlier regionally by −0.143 days/year and −0.363 days/year, respectively. Because of changes in SGS and EGS, length of growing season (LGS) across the GLRCA was shortened at a rate of −0.442 days/yr, which was mainly attributed to advanced EGS. Additionally, SGS of vegetation was negatively correlated with surface temperature but positively correlated with soil moisture and SPEI. These results indicated that surface temperature was a major determinant of advanced spring phenology, while increased soil moisture and mitigated drought would delay spring phenology. The response of autumn phenology to surface temperature and short-wave radiation varied across different climate zones. In arid climate zone, autumn phenology was obviously advanced with the increase of surface temperature and short-wave radiation. In cold climate zone, higher surface temperature and short-wave radiation postponed autumn phenology. Meanwhile, the thermal growing season did not accurately characterize the actual vegetation growing season because GLRCA phenology was different from most of Northern Hemisphere.

Influences of Shifted Vegetation Phenology on Runoff Across a Hydroclimatic Gradient.

Climate warming has changed vegetation phenology, and the phenology-associated impacts on terrestrial water fluxes remain largely unquantified. The impacts are linked to plant adjustments and responses to climate change and can be different in different hydroclimatic regions. Based on remote sensing data and observed river runoff of hydrological station from six river basins across a hydroclimatic gradient from northeast to southwest in China, the relative contributions of the vegetation (including spring and autumn phenology, growing season length (GSL), and gross primary productivity) and climatic factors affecting the river runoffs over 1982–2015 were investigated by applying gray relational analysis (GRA). We found that the average GSLs in humid regions (190–241 days) were longer than that in semi-humid regions (186–192 days), and the average GSLs were consistently extended by 4.8–13.9 days in 1982–2015 period in six river basins. The extensions were mainly linked to the delayed autumn phenology in the humid regions and to advanced spring phenology in the semi-humid regions. Across all river basins, the GRA results showed that precipitation (r = 0.74) and soil moisture (r = 0.73) determine the river runoffs, and the vegetation factors (VFs) especially the vegetation phenology also affected the river runoffs (spring phenology: r = 0.66; GSL: r = 0.61; autumn phenology: r = 0.59), even larger than the contribution from temperature (r = 0.57), but its relative importance is climatic region-dependent. Interestingly, the spring phenology is the main VF in the humid region for runoffs reduction, while both spring and autumn growth phenology are the main VFs in the semi-humid region, because large autumn phenology delay and less water supply capacity in spring amplify the effect of advanced spring phenology. This article reveals diverse linkages between climatic and VFs, and runoff in different hydroclimatic regions, and provides insights that vegetation phenology influences the ecohydrology process largely depending on the local hydroclimatic conditions, which improve our understanding of terrestrial hydrological responses to climate change.

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

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

Legacy effects of spring phenology on vegetation growth under preseason meteorological drought in the Northern Hemisphere

Revealing the physiological mechanisms and the implications of changes in spring phenology is crucial for understanding ecosystem productivity and functioning. Previous studies reported an increased vegetation growth associated with warming-induced earlier spring phenology, which can in turn make feedback to the climate systems and increase summer drought. The effects of preseason drought on spring phenology, especially their combined implications for vegetation growth, however, remain largely unknown. We first used four standard methods to extract dates for the start of spring phenology (SOS) from Normalized Difference Vegetation Index (NDVI3g) data across extratropical ecosystems in the Northern Hemisphere (>30°N) during 1982–2015, then calculated the sensitivity of SOS to preseason meteorological drought (preseason drought) using the Standardized Precipitation–Evapotranspiration Index (SPEI), and finally analyzed the coupling legacy effects of preseason drought and advanced SOS on seasonal vegetation growth across biomes. Our findings indicated that between 1982 and 2015 (1) SOS was positively correlated with preseason SPEI in 68.5% of pixels, 33.3% of which were significantly correlated (p < 0.05); the dominant timescales of preseason SPEI (preseason length) were 1 and 6 accumulated months. Mean SOS under extreme preseason drought also advanced in each biome. (2) Interestingly, SOS, rather than preseason drought or other climatic factors, had a dominant but opposite effect on spring (positively correlated) and summer (negatively correlated) vegetation growth. Most (86.4%) areas with increased spring growth were in regions with an earlier SOS, even under extreme preseason drought, but the combined effects of preseason drought and an earlier SOS greatly decreased summer vegetation growth in all biomes. Our results demonstrated that the preseason droughts during 1982–2015 had not yet reached the point beyond which they could overwhelm the role of warming in advancing spring phenology in most regions, but the earlier SOS induced by warming could exacerbate the effects of preseason drought and eventually have profound negative effects on vegetation growth in the middle and later stages of the growing season. This synthesis will advance our current understanding of mechanisms underlying atmosphere-biosphere feedbacks, and benefit biogeochemical models in simulating vegetation growth.

Contrasting temporal variations in responses of leaf unfolding to daytime and nighttime warming.

Earlier spring phenological events have been widely reported in plants under global warming. Recent studies reported a slowdown in the warming-induced advanced spring phenology in temperate regions. However, previous research mainly focused on daily mean temperature, thus neglecting the asymmetric phenological responses to daytime and nighttime temperature. Using long-term records of leaf unfolding in eight deciduous species at 1300sites across central Europe, we assessed and compared the effects of daytime temperature, nighttime temperature, and photoperiod on leaf unfolding during 1951-1980 and 1981-2013. Although leaf unfolding was advanced by daytime warming during 1951-2013, the advancing responses of leaf unfolding significantly decreased from 1951-1980 to 1981-2013 due to a lower accumulation of chilling units by daytime warming. Nighttime warming delayed leaf unfolding during 1951-1980 but advanced it during 1981-2013 due to a higher accumulation of chilling units by nighttime warming. In contrast, critical daylength and plasticity of leaf unfolding dates remained unchanged between 1951 and 2013. Our study provided evidence that daytime warming instead of nighttime warming accounts for the slowdown in the advancing spring phenology and implied that nighttime warming-induced earlier spring phenology may be buffering the slowdown of the advanced spring phenology by daytime warming. The response of spring phenology to nighttime temperature may override that to daytime temperature under the actual trends in global warming.

Widespread shifts in bird migration phenology are decoupled from parallel shifts in morphology.

Advancements in phenology and changes in morphology, including body size reductions, are among the most commonly described responses to globally warming temperatures. Although these dynamics are routinely explored independently, the relationships among them and how their interactions facilitate or constrain adaptation to climate change are poorly understood. In migratory species, advancing phenology may impose selection on morphological traits to increase migration speed. Advancing spring phenology might also expose species to cooler temperatures during the breeding season, potentially mitigating the effect of a warming global environment on body size. We use a dataset of birds that died after colliding with buildings in Chicago, IL to test whether changes in migration phenology are related to documented declines in body size and increases in wing length in 52 North American migratory bird species between 1978 and 2016. For each species, we estimate temporal trends in morphology and changes in the timing of migration. We then test for associations between species-specific rates of phenological and morphological changes while assessing the potential effects of migratory distance and breeding latitude. We show that spring migration through Chicago has advanced while the timing of fall migration has broadened as a result of early fall migrants advancing their migrations and late migrants delaying their migrations. Within species, we found that longer wing length was linked to earlier spring migration within years. However, we found no evidence that rates of phenological change across years, or migratory distance and breeding latitude, are predictive of rates of concurrent changes in morphological traits. These findings suggest that biotic responses to climate change are highly multidimensional and the extent to which those responses interact and influence adaptation to climate change requires careful examination.

Ecological responses to variation in seasonal snow cover.

Seasonal snow is among the most important factors governing the ecology of many terrestrial ecosystems, but rising global temperatures are changing snow regimes and driving widespread declines in the depth and duration of snow cover. Loss of the insulating snow layer will fundamentally change the environment. Understanding how individuals, populations, and communities respond to different snow conditions is thus essential for predicting and managing future ecosystem change. We synthesized 365 studies that examined ecological responses to variation in winter snow conditions. This research encompasses a broad range of methods (experimental manipulations, measurement of natural snow gradients, and long-term monitoring), locations (35 countries), study organisms (plants, mammals, arthropods, birds, fish, lichen, and fungi), and response measures. Earlier snowmelt was consistently associated with advanced spring phenology in plants, mammals, and arthropods. Reduced snow depth often increased mortality or physical injury in plants, although there were few clear effects on animals. Neither snow depth nor snowmelt timing had clear or consistent directional effects on body size of animals or biomass of plants. However, because 96% of studies were from the northern hemisphere, the generality of these trends across ecosystems and localities is also unclear. We identified substantial research gaps for several taxonomic groups and response types; research on wintertime responses was notably scarce. Future research should prioritize examination of the mechanisms underlying responses to changing snow conditions and the consequences of those responses for seasonally snow-covered ecosystems.

Spatial and temporal shifts in photoperiod with climate change.

Climate change causes both temporal (e.g. advancing spring phenology) and geographic (e.g. range expansion poleward) species shifts, which affect the photoperiod experienced at critical developmental stages ('experienced photoperiod'). As photoperiod is a common trigger of seasonal biological responses - affecting woody plant spring phenology in 87% of reviewed studies that manipulated photoperiod - shifts in experienced photoperiod may have important implications for future plant distributions and fitness. However, photoperiod has not been a focus of climate change forecasting to date, especially for early-season ('spring') events, often assumed to be driven by temperature. Synthesizing published studies, we find that impacts on experienced photoperiod from temporal shifts could be orders of magnitude larger than from spatial shifts (1.6h of change for expected temporal vs 1min for latitudinal shifts). Incorporating these effects into forecasts is possible by leveraging existing experimental data; we show that results from growth chamber experiments on woody plants often have data relevant for climate change impacts, and suggest that shifts in experienced photoperiod may increasingly constrain responses to additional warming. Further, combining modeling approaches and empirical work on when, where and how much photoperiod affects phenology could rapidly advance our understanding and predictions of future spatio-temporal shifts from climate change.

Climate warming prolongs the time interval between leaf‐out and flowering in temperate trees: Effects of chilling, forcing and photoperiod

Abstract Leaf‐out and flowering are two key phenological events of plants, denoting the respective onsets of visible vegetative growth and reproduction during the year. For each species, the schedule of vegetative growth and reproduction is crucial to the maximization of its fitness. Warming‐induced advances of leaf‐out and flowering have been reported frequently; however, it is unclear whether the responses of the two events are equal for any given species. Using long‐term phenological records in Europe, we examined simultaneously the responses of both leaf‐out and flowering of four common temperate tree species to climate warming and further examined the effects of winter chilling, spring forcing and photoperiod on the responses of the two events. We found that regardless whether flowering or leaf‐out occurred first, the first event advanced more than the second during 1950–2013, resulting in a prolonged time interval between the two events. The temporal changes were also supported by a similar geographical trend that the time interval between the two events increased from cold to warm sites. Due to the warming‐induced reduction in chilling, the spring forcing accumulated until the second event was increased more than the forcing accumulated until the first event, and that reduced the temperature sensitivity of the second event. In addition to the effect of chilling, the shorter photoperiod, associated with the advanced spring phenology, was also likely to substantially increase the spring forcing accumulated until the second event, which thus slowed down its advance, compared to the advance of the first event. The relative contributions of chilling and photoperiod to the increased forcing varied between species and events, with chilling mostly outweighing photoperiod. Synthesis. This study provides the large‐scale empirical evidence of prolonged time interval between leaf‐out and flowering with climate warming. The unequal advances of the two events may alter the partition of resources between vegetative growth and reproduction and cause different changes of spring frost damage to vegetative and reproductive tissues, which may alter species fitness and further affect ecosystem structure and function.

Response of barren-ground caribou to advancing spring phenology.

Phenological shifts are occurring in many ecosystems around the world. The capacity of species to adapt to changing phenology will be critical to their success under climate change scenarios. Failure to adjust migratory and reproductive timing to keep pace with the earlier onset of spring has led to negative demographic effects for populations of species across a variety of taxa. For caribou, there have been concerns that earlier spring green-up on calving areas might not be matched by earlier migration and parturition, potentially leading to a trophic mismatch with nutritional consequences for parturient and lactating caribou cows. However, there is limited evidence supporting these concerns. Here, we investigate the response of barren-ground caribou to changing spring phenology using data from telemetry and satellite imagery. From 2004 to 2016, we found that the average start of green-up on the calving area advanced by 7.25days, while the start of migration advanced by 13.64days, the end of migration advanced by 6.02days, and the date of peak calving advanced by 9.42days. Despite the advancing onset of green-up, we found no evidence for the development of a trophic mismatch because the advancing green-up coincided with earlier migration and calving by caribou. Changing snow cover on the late winter and migratory ranges was the most supported driver of advancing migratory behavior. The ability of caribou to adjust the timing of migratory and reproductive behavior in response to changing environmental conditions demonstrates the potential resilience of the species to some aspects of climate change.

Evolutionary Response to Climate Change in Migratory Pied Flycatchers.

Climate change is rapidly advancing spring phenology [1-3] but at different rates in different species [1, 4]. Whether these advances are solely driven by phenotypic plasticity [2, 5] or also involve evolution is hotly debated (e.g., [5-7]). In some species, including avian long-distance migrants, plastic responses to early springs may be constrained by inherited circannual timing programs [8, 9], making evolutionary adjustment theonly viable mechanism for keeping pace withshifting phenology [5, 10]. This constraint may becontributing to population declines in migratory species [5, 10-12]. To test whether a migrant's timing program has evolved [10, 12], wereplicated an experimental study of the annual cycle of long-distance migratory pied flycatchers (Ficedula hypoleuca) after 21 years of warming. Flycatchers are a model for studying constrained ecological responses to climate change [6, 10, 12, 13]. We show that the phase of the flycatcher circannual clock controlling spring moult, migration, and reproductive timing advanced by 9days. A nearby wild population mirrored these changes, concurrently advancing egg-laying by 11days. Furthermore, the time window during which wild flycatcher reproductive timing was most sensitive to ambient temperature advanced by 0.8days year-1. These results support a role of phenotypic evolution [14] in changing spring phenology [15, 16]. We suggest that the timing programs oflong-distance migratory birds may have greater adaptive potential than previously thought, leaving some scope for evolutionary rescue in a changing climate.

Cloud cover and delayed herbivory relative to timing of spring onset interact to dampen climate change impacts on net ecosystem exchange in a coastal Alaskan wetland

Rapid warming in northern ecosystems over the past four decades has resulted in earlier spring, increased precipitation, and altered timing of plant–animal interactions, such as herbivory. Advanced spring phenology can lead to longer growing seasons and increased carbon (C) uptake. Greater precipitation coincides with greater cloud cover possibly suppressing photosynthesis. Timing of herbivory relative to spring phenology influences plant biomass. None of these changes are mutually exclusive and their interactions could lead to unexpected consequences for Arctic ecosystem function. We examined the influence of advanced spring phenology, cloud cover, and timing of grazing on C exchange in the Yukon–Kuskokwim Delta of western Alaska for three years. We combined advancement of the growing season using passive-warming open-top chambers (OTC) with controlled timing of goose grazing (early, typical, and late season) and removal of grazing. We also monitored natural variation in incident sunlight to examine the C exchange consequences of these interacting forcings. We monitored net ecosystem exchange of C (NEE) hourly using an autochamber system. Data were used to construct daily light curves for each experimental plot and sunlight data coupled with a clear-sky model was used to quantify daily and seasonal NEE over a range of incident sunlight conditions. Cloudy days resulted in the largest suppression of NEE, reducing C uptake by approximately 2 g C m−2 d−1 regardless of the timing of the season or timing of grazing. Delaying grazing enhanced C uptake by approximately 3 g C m−2 d−1. Advancing spring phenology reduced C uptake by approximately 1.5 g C m−2 d−1, but only when plots were directly warmed by the OTCs; spring advancement did not have a long-term influence on NEE. Consequently, the two strongest drivers of NEE, cloud cover and grazing, can have opposing effects and thus future growing season NEE will depend on the magnitude of change in timing of grazing and incident sunlight.

Territory location and quality, together with climate, affect the timing of breeding in the white-throated dipper

Recent climate change has led to advanced spring phenology in many temperate regions. The phenological response to variation in the local environment, such as the habitat characteristics of the territories birds occupy, is less clear. The aim of this study is to understand how ecological conditions affect breeding time, and its consequences for reproduction, in a white-throated dipper Cinclus cinclus population in a river system in Norway during 34 years (1978–2011). Hatching date advanced almost nine days, indicating a response to higher temperatures and the advanced phenology in the area. Earlier breeding was found in warm springs and at lower altitudes. High population density facilitated earlier breeding close to the coast. Furthermore, when population density was low, breeding was early at territories that were rarely occupied, while in years with high density, breeding was early at territories that were frequently occupied. Also, when population density was low, earlier breeding occurred at territories that on average produced more offspring than other territories, while there was no difference in breeding time in high population years. Selection for early breeding was dependent on spring temperatures and high spring temperatures contributed to higher breeding success during the study period. We found that breeding phenology may have strong effects on fitness in the white-throated dipper, and thus that breeding time is an important ecological factor in a species that feeds mainly on aquatic rather than terrestrial prey.