Current issues in tropical phenology: a synthesis

We retrace the development of tropical phenology research, compare temperate phenology study to that in the tropics and highlight the advances currently being made in this flourishing discipline. The synthesis draws attention to how fundamentally different tropical phenology data can be to temperate data. Tropical plants lack a phase of winter dormancy and may grow and reproduce continually. Seasonal patterns in environmental parameters, such as rainfall, irradiance or temperature, do not necessarily coincide temporally, as they do in temperate climes. We review recent research on the drivers of phenophase cycles in individual trees, species and communities and highlight how significant innovations in biometric tools and approaches are being driven by the need to deal with circular data, the complexity of defining tropical seasons and the myriad growth and reproductive strategies used by tropical plants. We discuss how important the use of leaf phenology (or remotely‐sensed proxies of leaf phenophases) has become in tracking biome responses to climate change at the continental level and how important the phenophase of forests can be in determining local weather conditions. We also highlight how powerful analyses of plant responses are hampered at many tropical sites by a lack of contextual data on local environmental conditions. We conclude by arguing that there is a clear global benefit in increasing long term tropical phenology data collection and improving empirical collection of local climate measures, contemporary to the phenology data. Directing more resources to research in this sector will be widely beneficial.     

Delayed autumn phenology in the Northern Hemisphere is related to change in both climate and spring phenology

The timing of the end of the vegetation growing season (EOS) plays a key role in terrestrial ecosystem carbon and nutrient cycles. Autumn phenology is, however, still poorly understood, and previous studies generally focused on few species or were very limited in scale. In this study, we applied four methods to extract EOS dates from NDVI records between 1982 and 2011 for the Northern Hemisphere, and determined the temporal correlations between EOS and environmental factors (i.e., temperature, precipitation and insolation), as well as the correlation between spring and autumn phenology, using partial correlation analyses. Overall, we observed a trend toward later EOS in ~70% of the pixels in Northern Hemisphere, with a mean rate of 0.18 ± 0.38 days yr^?1. Warming preseason temperature was positively associated with the rate of EOS in most of our study area, except for arid/semi‐arid regions, where the precipitation sum played a dominant positive role. Interestingly, increased preseason insolation sum might also lead to a later date of EOS. In addition to the climatic effects on EOS, we found an influence of spring vegetation green‐up dates on EOS, albeit biome dependent. Our study, therefore, suggests that both environmental factors and spring phenology should be included in the modeling of EOS to improve the predictions of autumn phenology as well as our understanding of the global carbon and nutrient balances.     

Changes in autumn senescence in northern hemisphere deciduous trees: a meta-analysis of autumn phenology studies

Background and Aims Many individual studies have shown that the timing of leaf senescence in boreal and temperate deciduous forests in the northern hemisphere is influenced by rising temperatures, but there is limited consensus on the magnitude, direction and spatial extent of this relationship.Methods A meta-analysis was conducted of published studies from the peer-reviewed literature that reported autumn senescence dates for deciduous trees in the northern hemisphere, encompassing 64 publications with observations ranging from 1931 to 2010.Key Results Among the meteorological measurements examined, October temperatures were the strongest predictors of date of senescence, followed by cooling degree-days, latitude, photoperiod and, lastly, total monthly precipitation, although the strength of the relationships differed between high- and low-latitude sites. Autumn leaf senescence has been significantly more delayed at low (25° to 49°N) than high (50° to 70°N) latitudes across the northern hemisphere, with senescence across high-latitude sites more sensitive to the effects of photoperiod and low-latitude sites more sensitive to the effects of temperature. Delays in leaf senescence over time were stronger in North America compared with Europe and Asia.Conclusions The results indicate that leaf senescence has been delayed over time and in response to temperature, although low-latitude sites show significantly stronger delays in senescence over time than high-latitude sites. While temperature alone may be a reasonable predictor of the date of leaf senescence when examining a broad suite of sites, it is important to consider that temperature-induced changes in senescence at high-latitude sites are likely to be constrained by the influence of photoperiod. Ecosystem-level differences in the mechanisms that control the timing of leaf senescence may affect both plant community interactions and ecosystem carbon storage as global temperatures increase over the next century.     

Variable shifts in spring and autumn migration phenology in North American songbirds associated with climate change

Monitoring studies find that the timing of spring bird migration has advanced in recent decades, especially in Europe. Results for autumn migration have been mixed. Using data from Powdermill Nature Reserve, a banding station in western Pennsylvania, USA, we report an analysis of migratory timing in 78 songbird species from 1961 to 2006. Spring migration became significantly earlier over the 46-year period, and autumn migration showed no overall change. There was much variation among species in phenological change, especially in autumn. Change in timing was unrelated to summer range (local vs. northern breeders) or the number of broods per year, but autumn migration became earlier in neotropical migrants and later in short-distance migrants. The migratory period for many species lengthened because late phases of migration remained unchanged or grew later as early phases became earlier. There was a negative correlation between spring and autumn in long-term change, and this caused dramatic adjustments in the amount of time between migrations: the intermigratory periods of 10 species increased or decreased by > 15 days. Year-to-year changes in timing were correlated with local temperature (detrended) and, in autumn, with a regional climate index (detrended North Atlantic Oscillation). These results illustrate a complex and dynamic annual cycle in songbirds, with responses to climate change differing among species and migration seasons.     

Confounding effects of spatial variation on shifts in phenology

Shifts in the timing of life history events have become an important source of information about how organisms are responding to climate change. Phenological data have generally been treated as purely temporal, with scant attention to the inherent spatial aspects of such data. However, phenological data are tied to a specific location, and considerations of sampling design, both over space and through time, can critically affect the patterns that emerge. Focusing on flowering phenology, we describe how purely spatial shifts, such as adding new study plots, or the colonization of a study plot by a new species, can masquerade as temporal shifts. Such shifts can look like responses to climate change but are not. Furthermore, the same aggregate phenological curves can be composed of individuals with either very different or very similar phenologies. We conclude with a set of recommendations to avoid ambiguities arising from the spatiotemporal duality of phenological data.     

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.     

Direct and indirect effects of weather variability in a specialist butterfly

1. Climate change poses serious threats to the long‐term persistence of many animal and plant populations. Species having specific niche requirements, or characterised by highly co‐evolved interactions, will face the greatest challenges. An example is represented by Maculinea alcon (Denis & Schiffermüller), a monophagous and univoltine butterfly species, which lays eggs only on larval host plants which occur inparticular phenological conditions. 2. The present 2‐year study focused on two M. alcon populations, both located at the southern boundaries of the species, but facing different climatic conditions (360 m, low altitude versus 860 m, high altitude). Population vulnerability with respect to direct and indirect effects of climate change was analysed, focusing on two important aspects of butterfly biology, i.e. the flight activity of adults and the degree of synchrony in the larval plant–insect interactions. 3. It was observed that, when positive temperature anomalies are reached, the temperature can exert detrimental effects on adults' activity. At a low altitude, in a hotter than usual year, a temperature threshold was recorded (around 32 °C), above which the activity of butterflies is inhibited. In contrast, at a high altitude, temperature increases maintain the opportunity to enhance butterfly activity. Altitudinal differences were also observed in the phenology of the two interacting species, which generate stronger asynchrony at low altitudes. 4. High‐ and low‐altitude populations represent different conservation units: a global increase in temperature would pose a serious threat to the lowland populations, whereas high‐altitude populations would gain a greater role in assuring the persistence of M. alcon at its southern boundaries.     

Sessile oak forest plant community changes on the NE Iberian Peninsula over recent decades

AimsOur aims were 3-fold: (i) to determine whether global change has altered the composition and structure of the plant community found in the sessile oak forests on the NE Iberian Peninsula over the last decades, (ii) to establish whether the decline in forest exploitation activities that has taken place since the mid-20th century has had any effect on the forests and (iii) to ascertain whether there is any evidence of impact from climate warming.     

气候变化对鸟类影响的研究进展

气候变化对生物多样性的影响已成为热点问题.本文以鸟类为研究对象,根据鸟类受气候变化影响的最新研究成果,综述了气候变化对鸟类的分布、物候和种群等方面的影响.结果表明,在气候变化影响下,鸟类分布向高纬度或高海拔区移动,速度比以往加快,繁殖地和非繁殖地的分布移动变化并不相同,并且多数分布范围缩小,物候期发生复杂变化,种群数量下降明显.文章还讨论了该领域主要的预测和评估方法,以及进化适应等生物因素对气候变化预测结果的影响,除了以往单一的相关性模型外,目前应用最多的是集成模型,而未来最具发展潜力的是机理模型.进化适应方面的研究近来取得新进展,证实了生物个体积极应对气候变化影响的事实,从而对人为模型预测的准确性带来挑战.文章最后进行了总结和展望,结合国外研究经验和我国实际情况,提出一些建议:由于气候变化的影响及其研究是长期性的,从而对鸟类的历史监测数据提出很高的要求,当前我国急需建立一套长期、全面和可靠的鸟类数据监测系统;此外,人们需要综合评估现有各种预测模型的可靠性,在此基础上探索新的研究方法.     

Forecasting phenology: from species variability to community patterns

Shifts in species' phenology in response to climate change have wide-ranging consequences for ecological systems. However, significant variability in species' responses, together with limited data, frustrates efforts to forecast the consequences of ongoing phenological changes. Herein, we use a case study of three North American plant communities to explore the implications of variability across levels of organisation (within and among species, and among communities) for forecasting responses to climate change. We show how despite significant variation among species in sensitivities to climate, comparable patterns emerge at the community level once regional climate drivers are accounted for. However, communities differ with respect to projected patterns of divergence and overlap among their species' phenological distributions in response to climate change. These analyses and a review of hypotheses suggest how explicit consideration of spatial scale and levels of biological organisation may help to understand and forecast phenological responses to climate change.     

气候变化对植物-传粉昆虫的分布区和物候及其互作关系的影响

全球气候变化对生态系统的影响是人类社会面临的紧迫而又严峻的挑战。气候变化带来的极端气候事件的增多, 直接影响到生态系统生产力和服务功能。本文总结了气候变化对植物-传粉昆虫互作的研究进展, 强调植物-传粉昆虫互作网络结构和其时空演变的解析, 以及互作关系和功能性状重组研究的重要性。近年来在气温持续上升背景下对植物-传粉昆虫互作关系影响的研究也受到了更多关注, 这些研究主要集中在两方面: 一是植物和传粉昆虫分布区的变化, 包括部分种群可能灭绝; 二是物候的变化, 即植物花期和传粉昆虫活动期的改变。植物与传粉昆虫任何一方在空间或时间上的改变, 都会导致传粉关系的错配或丢失。此外, 也可能导致植物-传粉昆虫双方的功能性状及其耦合的改变, 从而影响其互作关系的稳定。建议在今后的研究中关注: (1)覆盖生物多样性的多个尺度的研究; (2)对植物-传粉者互作网络的长期监测; (3)重要指示物种繁殖适合度评价; (4)植物-传粉昆虫互作双方功能性状在时间和空间尺度上的变化, 及其互作关系的重组; (5)关键植物和传粉昆虫类群的评估和保护。     

Temperature,precipitation, and insolation effects on autumn vegetation phenology in temperate China

Autumn phenology plays a critical role in regulating climate–biosphere interactions. However, the climatic drivers of autumn phenology remain unclear. In this study, we applied four methods to estimate the date of the end of the growing season (EOS) across China's temperate biomes based on a 30‐year normalized difference vegetation index (NDVI) dataset from Global Inventory Modeling and Mapping Studies (GIMMS). We investigated the relationships of EOS with temperature, precipitation sum, and insolation sum over the preseason periods by computing temporal partial correlation coefficients. The results showed that the EOS date was delayed in temperate China by an average rate at 0.12 ± 0.01 days per year over the time period of 1982–2011. EOS of dry grassland in Inner Mongolia was advanced. Temporal trends of EOS determined across the four methods were similar in sign, but different in magnitude. Consistent with previous studies, we observed positive correlations between temperature and EOS. Interestingly, the sum of precipitation and insolation during the preseason was also associated with EOS, but their effects were biome dependent. For the forest biomes, except for evergreen needle‐leaf forests, the EOS dates were positively associated with insolation sum over the preseason, whereas for dry grassland, the precipitation over the preseason was more dominant. Our results confirmed the importance of temperature on phenological processes in autumn, and further suggested that both precipitation and insolation should be considered to improve the performance of autumn phenology models.     

Time of the season: the effect of host photoperiodism on diapause induction in an insect herbivore,Leptinotarsa decemlineata

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Using community photography to investigate phenology: A case study of coat molt in the mountain goat (Oreamnos americanus) with missing data

Participatory approaches, such as community photography, can engage the public in questions of societal and scientific interest while helping advance understanding of ecological patterns and processes. We combined data extracted from community‐sourced, spatially explicit photographs with research findings from 2018 fieldwork in the Yukon, Canada, to evaluate winter coat molt patterns and phenology in mountain goats (Oreamnos americanus), a cold‐adapted, alpine mammal. Leveraging the community science portals iNaturalist and CitSci, in less than a year we amassed a database of almost seven hundred unique photographs spanning some 4,500 km between latitudes 37.6°N and 61.1°N from 0 to 4,333 m elevation. Using statistical methods accounting for incomplete data, a common issue in community science datasets, we identified the effects of intrinsic (sex and presence of offspring) and broad environmental (latitude and elevation) factors on molt onset and rate and compared our findings with published data. Shedding occurred over a 3‐month period between 29 May and 6 September. Effects of sex and offspring on the timing of molt were consistent between the community‐sourced and our Yukon data and with findings on wild mountain goats at a long‐term research site in west‐central Alberta, Canada. Males molted first, followed by females without offspring (4.4 days later in the coarse‐grained, geographically wide community science sample; 29.2 days later in our fine‐grained Yukon sample) and lastly females with new kids (6.2; 21.2 days later, respectively). Shedding was later at higher elevations and faster at northern latitudes. Our findings establish a basis for employing community photography to examine broad‐scale questions about the timing of ecological events, as well as sex differences in response to possible climate drivers. In addition, community photography can help inspire public participation in environmental and outdoor activities specifically with reference to iconic wildlife.     

The seasonal timing of warming that controls onset of the growing season

Forecasting how global warming will affect onset of the growing season is essential for predicting terrestrial productivity, but suffers from conflicting evidence. We show that accurate estimates require ways to connect discrete observations of changing tree status (e.g., pre‐ vs. post budbreak) with continuous responses to fluctuating temperatures. By coherently synthesizing discrete observations with continuous responses to temperature variation, we accurately quantify how increasing temperature variation accelerates onset of growth. Application to warming experiments at two latitudes demonstrates that maximum responses to warming are concentrated in late winter, weeks ahead of the main budbreak period. Given that warming will not occur uniformly over the year, knowledge of when temperature variation has the most impact can guide prediction. Responses are large and heterogeneous, yet predictable. The approach has immediate application to forecasting effects of warming on growing season length, requiring only information that is readily available from weather stations and generated in climate models.     

The importance of phylogeny to the study of phenological response to global climate change

Climate change has resulted in major changes in the phenology—i.e. the timing of seasonal activities, such as flowering and bird migration—of some species but not others. These differential responses have been shown to result in ecological mismatches that can have negative fitness consequences. However, the ways in which climate change has shaped changes in biodiversity within and across communities are not well understood. Here, we build on our previous results that established a link between plant species'' phenological response to climate change and a phylogenetic bias in species'' decline in the eastern United States. We extend a similar approach to plant and bird communities in the United States and the UK that further demonstrates that climate change has differentially impacted species based on their phylogenetic relatedness and shared phenological responses. In plants, phenological responses to climate change are often shared among closely related species (i.e. clades), even between geographically disjunct communities. And in some cases, this has resulted in a phylogenetically biased pattern of non-native species success. In birds, the pattern of decline is phylogenetically biased but is not solely explained by phenological response, which suggests that other traits may better explain this pattern. These results illustrate the ways in which phylogenetic thinking can aid in making generalizations of practical importance and enhance efforts to predict species'' responses to future climate change.     

Predicting adaptation of phenology in response to climate change, an insect herbivore example

Climate change has led to an advance in phenology in many species. Synchrony in phenology between different species within a food chain may be disrupted if an increase in temperature affects the phenology of the different species differently, as is the case in the winter moth egg hatch–oak bud burst system. Operophtera brumata (winter moth) egg hatch date has advanced more than Quercus robur (pedunculate oak) bud burst date over the past two decades. Disrupted synchrony will lead to selection, and a response in phenology to this selection may lead to species genetically adapting to their changing environment. However, a prerequisite for such genetic change is that there is sufficient genetic variation and severe enough fitness consequences. So far, examples of observed genetic change have been few. Using a half-sib design, we demonstrate here that O. brumata egg-hatching reaction norm is heritable, and that genetic variation exists. Fitness consequences of even a few days difference between egg hatch and tree bud opening are severe, as we experimentally determined. Estimates of genetic variation and of fitness were then combined with a climate scenario to predict the rate and the amount of change in the eggs' response to temperature. We predict a rapid response to selection, leading to a restoration of synchrony of egg hatch with Q. robur bud opening. This study shows that in this case there is a clear potential to adapt – rapidly – to environmental change. The current observed asynchrony is therefore not due to a lack of genetic variation and at present it is unclear what is constraining O. brumata to adapt. This kind of model may be particularly useful in gaining insight in the predicted amount and rate of change due to environmental changes, given a certain genetic variation and selection pressure.     

Erosion of community diversity and stability by herbivore removal under warming

Climate change has the potential to influence the persistence of ecological communities by altering their stability properties. One of the major drivers of community stability is species diversity, which is itself expected to be altered by climate change in many systems. The extent to which climatic effects on community stability may be buffered by the influence of species interactions on diversity is, however, poorly understood because of a paucity of studies incorporating interactions between abiotic and biotic factors. Here, I report results of a 10-year field experiment, the past 7 years of which have focused on effects of ongoing warming and herbivore removal on diversity and stability within the plant community, where competitive species interactions are mediated by exploitation through herbivory. Across the entire plant community, stability increased with diversity, but both stability and diversity were reduced by herbivore removal, warming and their interaction. Within the most species-rich functional group in the community, forbs, warming reduced species diversity, and both warming and herbivore removal reduced the strength of the relationship between diversity and stability. Species interactions, such as exploitation, may thus buffer communities against destabilizing influences of climate change, and intact populations of large herbivores, in particular, may prove important in maintaining and promoting plant community diversity and stability in a changing climate.     

Larger temperature response of autumn leaf senescence than spring leaf‐out phenology

Climate warming is substantially shifting the leaf phenological events of plants, and thereby impacting on their individual fitness and also on the structure and functioning of ecosystems. Previous studies have largely focused on the climate impact on spring phenology, and to date the processes underlying leaf senescence and their associated environmental drivers remain poorly understood. In this study, experiments with temperature gradients imposed during the summer and autumn were conducted on saplings of European beech to explore the temperature responses of leaf senescence. An additional warming experiment during winter enabled us to assess the differences in temperature responses of spring leaf‐out and autumn leaf senescence. We found that warming significantly delayed the dates of leaf senescence both during summer and autumn warming, with similar temperature sensitivities (6–8 days delay per °C warming), suggesting that, in the absence of water and nutrient limitation, temperature may be a dominant factor controlling the leaf senescence in European beech. Interestingly, we found a significantly larger temperature response of autumn leaf senescence than of spring leaf‐out. This suggests a possible larger contribution of delays in autumn senescence, than of the advancement in spring leaf‐out, to extending the growing season under future warmer conditions.