全球变化下植物物候研究的关键问题

总结了全球变化下植物物候研究的主要进展,针对该领域国内外的几个热点问题进行了讨论。植物物候研究的重心从以前的野外观测和初步统计分析逐步过渡到以揭示物候周期的调控机制和环境效应为主,研究手段从植物物候对环境变化做出反应的表象描述转移到多尺度、多要素耦合关系的综合分析。随着学科交叉研究的不断深入,植物物候研究从植物个体及居群适应性研究转向植物物候变化对生态系统、气候演变、农业生产乃至人类健康等方面影响的系统评估。并且在该转变过程中出现了几个关键性问题,如不同温度带大气温度与光周期对植物物候期贡献力问题、植物物候变化对气候变暖的非线性响应特征、群落水平上植物物候研究的复杂性、以及农业生态系统中作物物候研究的重要性等。对我国植物物候研究现状和管理体系中亟待解决的问题提出了建议。     

Toward a synthetic understanding of the role of phenology in ecology and evolution

Phenology affects nearly all aspects of ecology and evolution. Virtually all biological phenomena—from individual physiology to interspecific relationships to global nutrient fluxes—have annual cycles and are influenced by the timing of abiotic events. Recent years have seen a surge of interest in this topic, as an increasing number of studies document phenological responses to climate change. Much recent research has addressed the genetic controls on phenology, modelling techniques and ecosystem-level and evolutionary consequences of phenological change. To date, however, these efforts have tended to proceed independently. Here, we bring together some of these disparate lines of inquiry to clarify vocabulary, facilitate comparisons among habitat types and promote the integration of ideas and methodologies across different disciplines and scales. We discuss the relationship between phenology and life history, the distinction between organismal- and population-level perspectives on phenology and the influence of phenology on evolutionary processes, communities and ecosystems. Future work should focus on linking ecological and physiological aspects of phenology, understanding the demographic effects of phenological change and explicitly accounting for seasonality and phenology in forecasts of ecological and evolutionary responses to climate change.     

A perspective on match/mismatch of phenology in community contexts

Climate change has significant impacts on phenology of various organisms in a species‐specific manner. Facing this problem, the match/mismatch hypothesis that phenological (a)synchrony with resource availability strongly influences recruitment success of a consumer population has recently received much attention. In this article, we discuss extending the conventional pairwise concept and demonstrate a community module‐based approach as an initial step for exploring community consequences of species‐specific phenological shifts caused by climate change. Our multispecies match/mismatch perspective leads to the prediction that phenological (a)synchrony among interacting species critically affects not only population recruitment of species but also key dynamical features of ecological communities such as trophic cascades, competitive hierarchies, and species coexistence. Explicit identification and consideration of species relationships is therefore desirable for a better understanding of seasonal community dynamics and thus community consequences of climate change‐induced phenological shifts.     

植物物候学研究进展

植物物候变化在研究陆地生态系统对气候变化的响应时被誉为"矿井中的金丝雀",全球气候变化愈演愈烈,重新引起了人们对植物物候研究的广泛关注。随着观测技术的发展,在各种空间和生态尺度上收集到的物候观测数据迅速累积,尽管已经在多个尺度上（物种、群落和景观尺度）观察到物候变化,但物候变化的机理仍然没有得到很好的理解。回顾了国内外植物物候研究的发展历程;总结了物候数据收集技术进展和全球物候变化的主要趋势;归纳了植物物候变化的机理与驱动因素;探讨了物候模型研究及物候对气候变化响应研究的主要方向。随着物候观测技术在不同尺度上应用的增加,物候研究进入了一个新的阶段。未来物候研究需要制定跨区域标准化观测指南,融合所有相关学科,改进物候模型,拓展研究区域;同时融合有效的历史物候资料,采用新技术和长期收集的物候数据为大数据时代植物物候学研究提供基础。     

Using phenological cameras to track the green up in a cerrado savanna and its on-the-ground validation

Plant phenology has gained new importance in the context of global change research, stimulating the development of novel technologies for phenological observations. Regular digital cameras have been effectively used as three-channel imaging sensors, providing measures of leaf color change or phenological shifts in plants. We monitored a species rich Brazilian cerrado savanna to assess the reliability of digital images to detect leaf-changing patterns. Analysis was conducted by extracting color information from selected parts of the image named regions of interest (ROIs). We aimed to answer the following questions: (i) Do digital cameras capture leaf changes in cerrado savanna vegetation? (ii) Can we detect differences in phenological changes among species crowns and the cerrado community? (iii) Is the greening pattern detected for each species by digital camera validated by our on-the-ground leafing phenology (direct observation of tree leaf changes)? We analyzed daily sequences of five images per hour, taken from 6:00 to 18:00 h, recorded during the cerrado main leaf flushing season. We defined 24 ROIs in the original digital image, including total or partial regions and crowns of six plant species. Our results indicated that: (i) for the studied period, single plant species ROIs were more sensitive to changes in relative green values than the community ROIs, (ii) three leaf strategies could be depicted from the species' ROI patterns of green color change, and (iii) the greening patterns and leaf functional groups were validated by our on-the-ground phenology. We concluded that digital cameras are reliable tools to monitor high diverse tropical seasonal vegetation and it is sensitive to inter-species differences of leafing patterns.     

Community‐wide changes in intertaxonomic temporal co‐occurrence resulting from phenological shifts

Global climate change is known to affect the assembly of ecological communities by altering species' spatial distribution patterns, but little is known about how climate change may affect community assembly by changing species' temporal co‐occurrence patterns, which is highly likely given the widely observed phenological shifts associated with climate change. Here, we analyzed a 29‐year phenological data set comprising community‐level information on the timing and span of temporal occurrence in 11 seasonally occurring animal taxon groups from 329 local meteorological observatories across China. We show that widespread shifts in phenology have resulted in community‐wide changes in the temporal overlap between taxa that are dominated by extensions, and that these changes are largely due to taxa's altered span of temporal occurrence rather than the degree of synchrony in phenological shifts. Importantly, our findings also suggest that climate change may have led to less phenological mismatch than generally presumed, and that the context under which to discuss the ecological consequences of phenological shifts should be expanded beyond asynchronous shifts.     

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.     

Climate change,uncertainty, and natural resource management

Climate change and its associated uncertainties are of concern to natural resource managers. Although aspects of climate change may be novel (e.g., system change and nonstationarity), natural resource managers have long dealt with uncertainties and have developed corresponding approaches to decision-making. Adaptive resource management is an application of structured decision-making for recurrent decision problems with uncertainty, focusing on management objectives, and the reduction of uncertainty over time. We identified 4 types of uncertainty that characterize problems in natural resource management. We examined ways in which climate change is expected to exacerbate these uncertainties, as well as potential approaches to dealing with them. As a case study, we examined North American waterfowl harvest management and considered problems anticipated to result from climate change and potential solutions. Despite challenges expected to accompany the use of adaptive resource management to address problems associated with climate change, we conclude that adaptive resource management approaches will be the methods of choice for managers trying to deal with the uncertainties of climate change. © 2010 The Wildlife Society.     

Managing biodiversity under climate change: challenges,frameworks, and tools for adaptation

The myriad challenges facing biodiversity under climate change are reflected in the challenges facing managers planning for these impacts. An ever-expanding number of recommendations and tools for climate change adaptation exist to aid managers in these efforts, yet navigating these various resources can lead to information overload and paralysis in decision-making. Here we provide a synthesis of the key considerations, approaches, and available tools for integrating climate change adaptation into management plans. We discuss principal elements in climate change adaptation—incorporating uncertainty through scenario planning and adaptive management—and review three leading frameworks for incorporating climate change adaptation into place-based biodiversity conservation planning. Finally, we identify the following key questions needed for long-term conservation planning and review the associated tools and techniques needed to address them: (1) How is the climate projected to change in my study area?; (2) How are non-climatic stressors projected to change?; (3) How vulnerable are species to climate change impacts?; (4) How are species ranges likely to respond?; and (5) How are management strategies expected to affect outcomes? In doing so, we aim to aid natural resource managers in navigating the burgeoning field of climate change adaptation planning and provide managers a roadmap for managing biodiversity under climate change.     

北京地区木本春季物候特征与生物学特性的关系

植物物候是植物生活史中的重要性状,也是指示气候与自然环境变化的重要指标,现已成为全球变化领域的研究热点之一。传统物候研究多假设物候由气候因素决定,如气温、降水、光照等,并主要从植物物候的年际变化角度探讨了气候因素对物候特征的影响。然而,不同物种的物候存在较大差异表明植物物候还与自身生物学特性（如系统发育和功能性状）有关,但植物生物学特性如何影响植物物候仍缺乏深入研究。基于北京地区44种木本植物1965-2018年的展叶始期和开花始期观测资料,以展叶始期和开花始期的3类物候特征（平均物候期、物候对温度的响应敏感度和物候期的积温需求）为例,探究植物物候特征与系统发育和功能性状的关系。首先,利用系统发育信号Blomberg’s K和进化模型检验植物物候特征是否具有系统发育保守性,并通过系统发育信号表征曲线直观表达植物物候特征的进化模式;之后,利用广义估计方程分析植物生活型、传粉型与物候特征的关系,以揭示不同植物的资源利用方式及生存策略的差异。研究发现：（1）除展叶始期的温度敏感度外,其余物候特征的进化均受随机遗传漂变和自然选择力的共同作用,可推断物候特征具有系统发育保守性,即亲缘关系越近的物种物候特征越相似。（2）开花始期的系统发育信号强度比展叶始期更大,表明繁殖物候的系统发育可能比生长物候更保守。（3）植物展叶始期及其积温需求与生活型密切相关。灌木比乔木的展叶时间早、积温需求少。植物开花始期与传粉型相关,风媒植物开花显著早于虫媒植物。研究成果有助于深入理解物候变化的生物学机制,对于丰富物候学的理论研究有重要意义,同时对植物保护也具有重要的指导价值。     

气候变化背景下橡胶树物候研究进展

橡胶树(Heveabrasiliensis)是广布于热带地区的经济林木，是战略物资天然橡胶的主要来源，其物候学的研究对胶园生产管理和评估热带地区植被对全球气候变化的响应方面具有重要意义。早期的物候研究主要服务于苗木繁育、割胶规划和抗逆栽培等生产应用；利用遥感监测植被物候日趋成熟，已广泛应用于橡胶树并成为主流的物候监测方法；橡胶树物候具有明显的时空异质性，对气候变化的响应较为复杂，其中温度和降水是关键影响因子，同时内因(品系、基因和树龄等)和外因(种植密度、地理位置和农业措施等)也共同影响了其物候。为更好服务天然橡胶产业的可持续发展和热区气候变化科学研究，未来的橡胶树物候研究应重点关注多源遥感数据的协同重建、物候指标提取算法的普适化和遥感预测模型的精准化。该文系统梳理了橡胶树物候的监测方法、服务价值、时空格局，提出了存在问题及未来研究方向。     

A systematic review of vegetation phenology in Africa

The study of vegetation phenology is important because it is a sensitive indicator of climate changes and it regulates carbon, energy and water fluxes between the land and atmosphere. Africa, which has 17% of the global forest cover, contributes significantly to the global carbon budget and has been identified as potentially highly vulnerable to climate change impacts. In spite of this, very little is known about vegetation phenology across Africa and the factors regulating vegetation growth and dynamics. Hence, this review aimed to provide a synthesis of studies of related Africa's vegetation phenology and classify them based on the methods and techniques used in order to identify major research gaps. Significant increases in the number of phenological studies in the last decade were observed, with over 70% of studies adopting a satellite-based remote sensing approach to monitor vegetation phenology. Whereas ground based studies that provide detailed characterisation of vegetation phenological development, occurred rarely in the continent. Similarly, less than 14% of satellite-based remote sensing studies evaluated vegetation phenology at the continental scale using coarse spatial resolution datasets. Even more evident was the lack of research focusing on the impacts of climate change on vegetation phenology. Consequently, given the importance and the uniqueness of both methods of phenological assessment, there is need for more ground-based studies to enable greater understanding of phenology at the species level. Likewise, finer spatial resolution satellite sensor data for regional phenological assessment is required, with a greater focus on the relationship between climate change and vegetation phenological changes. This would contribute greatly to debates over climate change impacts and, most importantly, climate change mitigation strategies.     

Preparing Wildlife for Climate Change: How Far Have We Come?

Global biodiversity is in unprecedented decline and on-the-ground solutions are imperative for conservation. Although there is a large volume of evidence related to climate change effects on wildlife, research on climate adaptation strategies is lagging. To assess the current state of knowledge in climate adaptation, we conducted a comprehensive literature review and evaluated 1,346 peer-reviewed publications for management recommendations designed to address the consequences of climate change on wildlife populations. From 509 publications, we identified 2,306 recommendations and employed both qualitative and quantitative methods for data analysis. Although we found an increase in the volume and diversity of recommendations since 2007, a focus on protected areas (26%, 596 of 2,306 recommendations) and the non-reserve matrix (12%, 276 of 2,306 recommendations) remained prominent in the climate adaptation literature. Common concepts include protected areas, invasive species, ecosystem services, adaptive management, stepping stones, assisted migration, and conservation easements. In contrast, only 1% of recommendations focused on reproduction (n = 26), survival (n = 14), disease (n = 26), or human-wildlife conflict (n = 24). Few recommendations reflected the potential for local-scale management interventions. We demonstrate limited advancement in preparing natural resource managers in climate adaptation at local, management-relevant scales. Additional research is needed to identify and evaluate climate adaptation strategies aimed at reducing the vulnerability of wildlife to contemporary climate change. © 2020 The Wildlife Society.     

Phenological plasticity is a poor predictor of subalpine plant population performance following experimental climate change

Phenological shifts, changes in the seasonal timing of life cycle events, are among the best documented responses of species to climate change. However, the consequences of these phenological shifts for population dynamics remain unclear. Population growth could be enhanced if species that advance their phenology benefit from longer growing seasons and gain a pre-emptive advantage in resource competition. However, it might also be reduced if phenological advances increase exposure to stresses, such as herbivores and, in colder climates, harsh abiotic conditions early in the growing season. We exposed subalpine grasslands to ~3 K of warming by transplanting intact turfs from 2000 m to 1400 m elevation in the eastern Swiss Alps, with turfs transplanted within the 2000 m site acting as a control. In the first growing season after transplantation, we recorded species’ flowering phenology at both elevations. We also measured species’ cover change for three consecutive years as a measure of plant performance. We used models to estimate species’ phenological plasticity (the response of flowering time to the change in climate) and analysed its relationship with cover changes following climate change. The phenological plasticity of the 18 species in our study varied widely but was unrelated to their changes in cover. Moreover, early- and late-flowering species did not differ in their cover response to warming, nor in the relationship between cover changes and phenological plasticity. These results were replicated in a similar transplant experiment within the same subalpine community, established one year earlier and using larger turfs. We discuss the various ecological processes that can be affected by phenological shifts, and argue why the population-level consequences of these shifts are likely to be species- and context-specific. Our results highlight the importance of testing assumptions about how warming-induced changes in phenotypic traits, like phenology, impact population dynamics.     

A framework for assessing the vulnerability of species to climate change: a case study of the Australian elapid snakes

Frameworks that provide a system for assessing species according to their vulnerability to climate change can offer considerable guidance to conservation managers who need to allocate limited resources among a large number of taxa. To date, climate change vulnerability assessments have largely been based on projected changes in range size derived from the output of species distribution models (SDMs). A criticism of risk assessments based solely on these models is that information on species ecological and life history traits is lacking. Accordingly, we developed a points-based framework for assessing species vulnerability to climate change that considered species traits together with the projections of SDMs. Applying this method to the Australian elapid snakes (family Elapidae), we determined which species may be particularly susceptible in the future and assessed broad-scale biogeographic patterns in species vulnerability. By offering a more comprehensive and rigorous method for assessing vulnerability than those based solely on SDMs, this framework provides greater justification for resource allocation, and can help guide decisions regarding the most appropriate adaptation strategies.     

Environmental controls on the phenology of moths: predicting plasticity and constraint under climate change

Ecological systems have naturally high interannual variance in phenology. Component species have presumably evolved to maintain appropriate phenologies under historical climates, but cases of inappropriate phenology can be expected with climate change. Understanding controls on phenology permits predictions of ecological responses to climate change. We studied phenological control systems in Lepidoptera by analyzing flight times recorded at a network of sites in Finland. We evaluated the strength and form of controls from temperature and photoperiod, and tested for geographic variation within species. Temperature controls on phenology were evident in 51% of 112 study species and for a third of those thermal controls appear to be modified by photoperiodic cues. For 24% of the total, photoperiod by itself emerged as the most likely control system. Species with thermal control alone should be most immediately responsive in phenology to climate warming, but variably so depending upon the minimum temperature at which appreciable development occurs and the thermal responsiveness of development rate. Photoperiodic modification of thermal controls constrains phenotypic responses in phenologies to climate change, but can evolve to permit local adaptation. Our results suggest that climate change will alter the phenological structure of the Finnish Lepidoptera community in ways that are predictable with knowledge of the proximate physiological controls. Understanding how phenological controls in Lepidoptera compare to that of their host plants and enemies could permit general inferences regarding climatic effects on mid- to high-latitude ecosystems.     

Macrophenology: insights into the broad-scale patterns,drivers, and consequences of phenology

Plant phenology research has surged in recent decades, in part due to interest in phenological sensitivity to climate change and the vital role phenology plays in ecology. Many local-scale studies have generated important findings regarding the physiology, responses, and risks associated with shifts in plant phenology. By comparison, our understanding of regional- and global-scale phenology has been largely limited to remote sensing of green-up without the ability to differentiate among plant species. However, a new generation of analytical tools and data sources—including enhanced remote sensing products, digitized herbarium specimen data, and public participation in science—now permits investigating patterns and drivers of phenology across extensive taxonomic, temporal, and spatial scales, in an emerging field that we call macrophenology. Recent studies have highlighted how phenology affects dynamics at broad scales, including species interactions and ranges, carbon fluxes, and climate. At the cusp of this developing field of study, we review the theoretical and practical advances in four primary areas of plant macrophenology: (1) global patterns and shifts in plant phenology, (2) within-species changes in phenology as they mediate species' range limits and invasions at the regional scale, (3) broad-scale variation in phenology among species leading to ecological mismatches, and (4) interactions between phenology and global ecosystem processes. To stimulate future research, we describe opportunities for macrophenology to address grand challenges in each of these research areas, as well as recently available data sources that enhance and enable macrophenology research.     

The eco‐evolutionary consequences of interspecific phenological asynchrony – a theoretical perspective

The timing of biological events (phenology) is an important aspect of both a species’ life cycle and how it interacts with other species and its environment. Patterns of phenological change have been given much scientific attention, particularly recently in relation to climate change. For pairs of interacting species, if their rates of phenological change differ, then this may lead to asynchrony between them and disruption of their ecological interactions. However it is often difficult to interpret differential rates of phenological change and to predict their ecological and evolutionary consequences. We review theoretical results regarding this topic, with special emphasis on those arising from life history theory, evolutionary game theory and population dynamic models. Much ecological research on phenological change builds upon the concept of match/mismatch, so we start by putting forward a simple but general model that captures essential elements of this concept. We then systematically compare the predictions of this baseline model with expectations from theory in which additional ecological mechanisms and features of species life cycles are taken into account. We discuss the ways in which the fitness consequences of interspecific phenological asynchrony may be weak, strong, or idiosyncratic. We discuss theory showing that synchrony is not necessarily an expected evolutionary outcome, and how population densities are not necessarily maximized by adaptation, and the implications of these findings. By bringing together theoretical developments regarding the eco‐evolutionary consequences of phenological asynchrony, we provide an overview of available alternative hypotheses for interpreting empirical patterns as well as the starting point for the next generation of theory in this field.