Marine biological shifts and climate

Phenological, biogeographic and community shifts are among the reported responses of marine ecosystems and their species to climate change. However, despite both the profound consequences for ecosystem functioning and services, our understanding of the root causes underlying these biological changes remains rudimentary. Here, we show that a significant proportion of the responses of species and communities to climate change are deterministic at some emergent spatio-temporal scales, enabling testable predictions and more accurate projections of future changes. We propose a theory based on the concept of the ecological niche to connect phenological, biogeographic and long-term community shifts. The theory explains approximately 70% of the phenological and biogeographic shifts of a key zooplankton Calanus finmarchicus in the North Atlantic and approximately 56% of the long-term shifts in copepods observed in the North Sea during the period 1958–2009.     

Climate variability and the timing of spring raptor migration in eastern North America

Many birds have advanced their spring migration and breeding phenology in response to climate change, yet some long‐distance migrants appear constrained in their adjustments. In addition, bird species with long generation times and those in higher trophic positions may also be less able to track climate‐induced shifts in food availability. Migratory birds of prey may therefore be particularly vulnerable to climate change because: 1) most are long‐lived and have relatively low reproductive capacity, 2) many feed predominately on insectivorous passerines, and 3) several undertake annual migrations totaling tens of thousands of kilometers. Using multi‐decadal datasets for 14 raptor species observed at six sites across the Great Lakes region of North America, we detected phenological shifts in spring migration consistent with decadal climatic oscillations and global climate change. While the North Atlantic and El Niño Southern Oscillations exerted heterogeneous effects on the phenology of a few species, arrival dates more generally advanced by 1.18 d per decade, a pattern consistent with the effects of global climate change. After accounting for heterogeneity across observation sites, five of the 10 most abundant species advanced the bulk of their spring migration phenology. Contrary to expectations, we found that long‐distance migrants and birds with longer generation times tended to make the greatest advancements to their spring migration. Such results may indicate that phenotypic plasticity can facilitate climatic responses among these long‐lived predators.     

Horizontal distribution of microprotist community structure in the western Arctic Ocean during late summer and early fall of 2010

The western Arctic Ocean is composed of two regions: the southern shelf and the northern basin, whereas the marine ecosystem structure is expected to vary between the regions, little information is available, particularly for the planktonic protist community. In this study, we surveyed the horizontal distribution of microprotists (diatoms, dinoflagellates and ciliates) at 59 stations in the western Arctic Ocean during September and October of 2010. The abundances of diatoms, dinoflagellates and ciliates were 0–138,640, 0–16,460 and 0–10,933 cells L^?1, respectively, and all of the abundances were higher on the Chukchi Sea shelf. Cluster analysis based on abundance separated the microprotist community into five groups, which contain 25, 22, 6, 4 and 2 stations. The largest group was observed on the Chukchi Sea shelf, showing a high abundance predominated by diatoms (78 % of total abundance). The second group was observed from the East Siberian Sea to the Canada Basin, characterised by low abundance and ciliate dominance (36 % of total abundance). Because of the high abundance and predominance of diatoms, the former group is characterised by eutrophic waters, which are enhanced by the continuous inflow of the nutrient-rich Pacific Water through the Bering Strait. Due to the low abundance and the dominance of ciliates, the latter group is dominated by organisms of the microbial food web. The remaining three groups were smaller and located between the two large groups. The distribution of these three groups may be based on complex physical structures, such as the anticyclonic eddy near the shelf break.     

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.     

Leaf phenology in 22 North American tree species during the 21st century

Recent shifts in phenology are the best documented biological response to current anthropogenic climate change, yet remain poorly understood from a functional point of view. Prevailing analyses are phenomenological and approximate, only correlating temperature records to imprecise records of phenological events. To advance our understanding of phenological responses to climate change, we developed, calibrated, and validated process-based models of leaf unfolding for 22 North American tree species. Using daily meteorological data predicted by two scenarios (A2: +3.2 °C and B2: +1 °C) from the HadCM3 GCM, we predicted and compared range-wide shifts of leaf unfolding in the 20th and 21st centuries for each species. Model predictions suggest that climate change will affect leaf phenology in almost all species studied, with an average advancement during the 21st century of 5.0 days in the A2 scenario and 9.2 days in the B2 scenario. Our model also suggests that lack of sufficient chilling temperatures to break bud dormancy will decrease the rate of advancement in leaf unfolding date during the 21st century for many species. Some temperate species may even have years with abnormal budburst due to insufficient chilling. Species fell into two groups based on their sensitivity to climate change: (1) species that consistently had a greater advance in their leaf unfolding date with increasing latitude and (2) species in which the advance in leaf unfolding differed from the center to the northern vs. southern margins of their range. At the interspecific level, we predicted that early-leafing species tended to show a greater advance in leaf unfolding date than late-leafing species; and that species with larger ranges tend to show stronger phenological changes. These predicted changes in phenology have significant implications for the frost susceptibility of species, their interspecific relationships, and their distributional shifts.     

Phenological indices of avian reproduction: cryptic shifts and prediction across large spatial and temporal scales

Climate change‐induced shifts in phenology have important demographic consequences, and are frequently used to assess species' sensitivity to climate change. Therefore, developing accurate phenological predictions is an important step in modeling species' responses to climate change. The ability of such phenological models to predict effects at larger spatial and temporal scales has rarely been assessed. It is also not clear whether the most frequently used phenological index, namely the average date of a phenological event across a population, adequately captures phenological shifts in the distribution of events across the season. We use the long‐tailed tit Aegithalos caudatus (Fig. 1) as a case study to explore these issues. We use an intensive 17‐year local study to model mean breeding date and test the capacity of this local model to predict phenology at larger spatial and temporal scales. We assess whether local models of breeding initiation, termination, and renesting reveal phenological shifts and responses to climate not detected by a standard phenological index, that is, population average lay date. These models take predation timing/intensity into account. The locally‐derived model performs well at predicting phenology at the national scale over several decades, at both high and low temperatures. In the local model, a trend toward warmer Aprils is associated with a significant advance in termination dates, probably in response to phenological shifts in food supply. This results in a 33% reduction in breeding season length over 17 years – a substantial loss of reproductive opportunity that is not detected by the index of population average lay date. We show that standard phenological indices can fail to detect patterns indicative of negative climatic effects, potentially biasing assessments of species' vulnerability to climate change. More positively, we demonstrate the potential of detailed local studies for developing broader‐scale predictive models of future phenological shifts.     

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。     

Identifying potential evolutionary consequences of climate-driven phenological shifts

Climate change is shifting the phenology of many species throughout the world. While the interspecific consequences of these phenological shifts have been well documented, the intraspecific shifts and their resultant evolutionary consequences remain relatively unexplored. Here, we present a conceptual framework and overview of how phenological shifts within species can drive evolutionary change. We suggest that because the impacts of climate change are likely to vary across the range of a species and differentially impact individuals, phenological shifts may often be highly variable both within and among populations. Together these changes have the potential to alter existing patterns of gene flow and influence evolutionary trajectories by increasing phenological isolation and connectivity. Recent research examining the response of species to contemporary climate change suggests that both phenological isolation and connectivity may be likely responses to future climate change. However, recent studies also show mixed results on whether adaptive responses to climate change are likely to occur, as some populations have already shown adaptive responses to changing climate, while others have not despite fitness costs. While predicting the exact consequences of intraspecific phenological shifts may be difficult, identifying the evolutionary implications of these shifts will allow a better understanding of the effects of future climate change on species persistence and adaptation.     

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。     

Phaeocystis colony distribution in the North Atlantic Ocean since 1948, and interpretation of long-term changes in the Phaeocystis hotspot in the North Sea

Monitoring of Phaeocystis since 1948 during the Continuous Plankton Recorder survey indicates that over the last 5.5 decades the distribution of its colonies in the North Atlantic Ocean was not restricted to neritic waters: occurrence was also recorded in the open Atlantic regions sampled, most frequently in the spring. Apparently, environmental conditions in open ocean waters, also those far offshore, are suitable for complete lifecycle development of colonies (the only stage recorded in the survey). In the North Sea the frequency of occurrence was also highest in spring. Its southeastern part was the Phaeocystis abundance hotspot of the whole area covered by the survey. Frequency was especially high before the 1960s and after the 1980s, i.e., in the periods when anthropogenic nutrient enrichment was relatively low. Changes in eutrophication have obviously not been a major cause of long-term Phaeocystis variation in the southeastern North Sea, where total phytoplankton biomass was related significantly to river discharge. Evidence is presented for the suggestion that Phaeocystis abundance in the southern North Sea is to a large extent determined by the amount of Atlantic Ocean water flushed in through the Dover Strait. Since Phaeocystis plays a key role in element fluxes relevant to climate the results presented here have implications for biogeochemical models of cycling of carbon and sulphur. Sea-to-air exchange of CO₂ and dimethyl sulphide (DMS) has been calculated on the basis of measurements during single-year cruises. The considerable annual variation in phytoplankton and in its Phaeocystis component reported here does not warrant extrapolation of such figures.     

Scale-dependent climate signals drive breeding phenology of three seabird species

Breeding at the right time is essential for animals in seasonal climates in order to ensure that the energy demands of reproduction, particularly the nutritional requirements of growing young, coincide with peak food availability. Global climate change is likely to cause shifts in the timing of peak food availability, and in order to adapt successfully to current and future climate change, animals need to be able to adjust the time at which they initiate breeding. Many animals use environmental cues available before the breeding season to predict the seasonal peak in food availability and adjust their phenology accordingly. We tested the hypothesis that regulation of breeding onset should reflect the scale at which organisms perceive their environment by comparing phenology of three seabird species at a North Sea colony. As predicted, the phenology of two dispersive species, black-legged kittiwake ( Rissa tridactyla ) and common guillemot ( Uria aalge ), correlated with a large-scale environmental cue (the North Atlantic Oscillation), whereas a resident species, European shag ( Phalacrocorax aristotelis ), was more affected by local conditions (sea surface temperature) around the colony. Annual mean breeding success was lower in late years for European shags, but not for the other two species. Since correlations among climate patterns at different scales are likely to change in the future, these findings have important implications for how migratory animals can respond to future climate change.     

Diel feeding pattern and prey selection of mesozooplankton on microplankton community

Mesozooplankton play an important role in transporting energy from microphytoplankton and microzooplankton to higher trophic levels. However there were few studies on the diel feeding patterns and prey selectivity of mesozooplankton. We conducted feeding experiments of mesozooplankton in the East China Sea to determine their respective diel feeding patterns on diatoms, ciliates and dinoflagellates, and to assess the contribution of these prey items to mesozooplankton diet. The results showed higher mesozooplankton grazing rates on ciliates and dinoflagellates than on diatoms at the day time, and the opposite pattern at the night time. A significant prey selection was observed, in which mesozooplankton positively selected ciliates and dinoflagellates during day and diatoms at night. The different grazing reactions of mesozooplankton toward each prey item might be related to the mobility of the prey. In all, microzooplankton (ciliates and dinoflagellates) provided the majority of the mesozooplankton carbon ingestion, even at a station dominated by small pennate diatoms. In particular, dinoflagellates are an important prey of mesozooplankton in the East China Sea and their contribution to the diet of mesozooplankton is unproportionate to their abundance.     

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.     

Seasonality of North Atlantic phytoplankton from space: impact of environmental forcing on a changing phenology (1998–2012)

Seasonal pulses of phytoplankton drive seasonal cycles of carbon fixation and particle sedimentation, and might condition recruitment success in many exploited species. Taking advantage of long‐term series of remotely sensed chlorophyll a (1998–2012), we analyzed changes in phytoplankton seasonality in the North Atlantic Ocean. Phytoplankton phenology was analyzed based on a probabilistic characterization of bloom incidence. This approach allowed us to detect changes in the prevalence of different seasonal cycles and, at the same time, to estimate bloom timing and magnitude taking into account uncertainty in bloom detection. Deviations between different sensors stressed the importance of a prolonged overlap between successive missions to ensure a correct assessment of phenological changes, as well as the advantage of semi‐analytical chlorophyll algorithms over empirical ones to reduce biases. Earlier and more intense blooms were detected in the subpolar Atlantic, while advanced blooms of less magnitude were common in the Subtropical gyre. In the temperate North Atlantic, spring blooms advanced their timing and decreased in magnitude, whereas fall blooms delayed and increased their intensity. At the same time, the prevalence of locations with a single autumn/winter bloom or with a bimodal seasonal cycle increased, in consonance with a poleward expansion of subtropical conditions. Changes in bloom timing and magnitude presented a clear signature of environmental factors, especially wind forcing, although changes on incident photosynthetically active radiation and sea surface temperature were also important depending on latitude. Trends in bloom magnitude matched changes in mean chlorophyll a during the study period, suggesting that seasonal peaks drive long‐term trends in chlorophyll a concentration. Our results link changes in North Atlantic climate with recent trends in the phenology of phytoplankton, suggesting an intensification of these impacts in the near future.     

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.     

Effects of climate-driven temperature changes on the diversity of freshwater macroinvertebrates

Increasing temperatures due to climate change were found to influence abundance and timing of species in numerous ways. Whereas many studies have investigated climate-induced effects on the phenology and abundance of single species, less is known about climate-driven shifts in the diversity and composition of entire communities. Analyses of long-term data sets provide the potential to reveal such relationships. We analysed time series of entire communities of macrozoobenthos in lakes and streams in Northern Europe. There were no direct linear effects of temperature and climate indices (North Atlantic Oscillation index) on species composition and diversity, but using multivariate statistics we were able to show that trends in average temperature have already had profound impacts on species composition in lakes. These significant temperature signals on species composition were evident even though we analysed comparatively short time periods of 10–15 years. Future climate shifts may thus induce strong variance in community composition. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users. Priority programme of the German Research Foundation—contribution 6.     

Tell me what you eat and I'll tell you when you fly: diet can predict phenological changes in response to climate change

Changes in phenology are correlated with climate change. However, we still struggle to understand the traits making species susceptible to climate change, and the implications of species' reactions for communities and food webs. Butterflies and moths are an ecologically important group that have shown pronounced phenological changes over the last decades. Tests using a > 150-year dataset from 566 European butterfly and moth species demonstrated that variation in phenological change was strongly related to traits describing plant-herbivore interactions (larval diet breadth, diet composition), and the life cycle. The results indicate that climate change related shifts in phenology are correlated with the seasonal availability and palatability of food plants. Lepidopterans feeding on herbaceous plants showed smaller shifts in flight periods but larger increases in voltinism than lepidopterans feeding on woody plants. Consequently, the effect of herbivorous lepidopterans may increase in herb-rich grassland ecosystems under warmer conditions, and not in forest ecosystems.     

Climate change and oceanic barriers: genetic differentiation in Pomatomus saltatrix (Pisces: Pomatomidae) in the North Atlantic Ocean and the Mediterranean Sea

Nucleotide variation of partial cytochrome b sequences was analysed in the bluefish Pomatomus saltatrix to investigate the population-structuring roles of climate change and oceanic barriers. Western and eastern North Atlantic Ocean populations appeared to be totally isolated, with the latter connected to the Mediterranean Sea within which further structuring occurred.     

Mismatch managed? Phenological phase extension as a strategy to manage phenological asynchrony in plant–animal mutualisms

Species‐specific shifts in phenology (timing of periodic life cycle events) are occurring with climate change and are already disrupting interactions within and among trophic levels. Phenological phase duration (e.g. beginning to end of flowering) and complementarity (patterns of nonoverlap), and their responses to changing conditions, will be important determinants of species' adaptive capacity to these shifts. Evidence indicates that extension of phenological duration of mutualistic partners could buffer negative impacts that occur with phenological shifts. Therefore, we suggest that techniques to extend the length of phenological duration will contribute to management of systems experiencing phenological asynchrony. Techniques of phenological phase extension discussed include the role of abiotic heterogeneity, genetic and species diversity, and alteration of population timing. We explore these approaches with the goal of creating a framework to build adaptive capacity and address phenological asynchrony in plant–animal mutualisms under climate change.     

Impact of climate change on plant phenology in Mediterranean ecosystems

Plant phenology is strongly controlled by climate and has consequently become one of the most reliable bioindicators of ongoing climate change. We used a dataset of more than 200 000 records for six phenological events of 29 perennial plant species monitored from 1943 to 2003 for a comprehensive assessment of plant phenological responses to climate change in the Mediterranean region. Temperature, precipitation and North Atlantic Oscillation (NAO) were studied together during a complete annual cycle before phenological events to determine their relative importance and potential seasonal carry‐over effects. Warm and dry springs under a positive phase of NAO advance flowering, leaf unfolding and fruiting dates and lengthen the growing season. Spatial variability of dates (range among sites) was also reduced during warm and dry years, especially for spring events. Climate during previous weeks to phenophases occurrence had the greatest impact on plants, although all events were also affected by climate conditions several months before. Immediate along with delayed climate effects suggest dual triggers in plant phenology. Climatic models accounted for more than 80% of variability in flowering and leaf unfolding dates, and in length of the growing season, but for lower proportions in fruiting and leaf falling. Most part of year‐to‐year changes in dates was accounted for temperature, while precipitation and NAO accounted for <10% of dates' variability. In the case of flowering, insect‐pollinated species were better modelled by climate than wind‐pollinated species. Differences in temporal responses of plant phenology to recent climate change are due to differences in the sensitivity to climate among events and species. Spring events are changing more than autumn events as they are more sensitive to climate and are also undergoing the greatest alterations of climate relative to other seasons. In conclusion, climate change has shifted plant phenology in the Mediterranean region.