Temperature alone does not explain phenological variation of diverse temperate plants under experimental warming

Anthropogenic climate change has altered temperate forest phenology, but how these trends will play out in the future is controversial. We measured the effect of experimental warming of 0.6–5.0 °C on the phenology of a diverse suite of 11 plant species in the deciduous forest understory (Duke Forest, North Carolina, USA) in a relatively warm year (2011) and a colder year (2013). Our primary goal was to dissect how temperature affects timing of spring budburst, flowering, and autumn leaf coloring for functional groups with different growth habits, phenological niches, and xylem anatomy. Warming advanced budburst of six deciduous woody species by 5–15 days and delayed leaf coloring by 18–21 days, resulting in an extension of the growing season by as much as 20–29 days. Spring temperature accumulation was strongly correlated with budburst date, but temperature alone cannot explain the diverse budburst responses observed among plant functional types. Ring‐porous trees showed a consistent temperature response pattern across years, suggesting these species are sensitive to photoperiod. Conversely, diffuse‐porous species responded differently between years, suggesting winter chilling may be more important in regulating budburst. Budburst of the ring‐porous Quercus alba responded nonlinearly to warming, suggesting evolutionary constraints may limit changes in phenology, and therefore productivity, in the future. Warming caused a divergence in flowering times among species in the forest community, resulting in a longer flowering season by 10‐16 days. Temperature was a good predictor of flowering for only four of the seven species studied here. Observations of interannual temperature variability overpredicted flowering responses in spring‐blooming species, relative to our warming experiment, and did not consistently predict even the direction of flowering shifts. Experiments that push temperatures beyond historic variation are indispensable for improving predictions of future changes in phenology.     

Phylogenetic conservatism and trait correlates of spring phenological responses to climate change in northeast China

Climate change has resulted in major changes in plant phenology across the globe that includes leaf‐out date and flowering time. The ability of species to respond to climate change, in part, depends on their response to climate as a phenological cue in general. Species that are not phenologically responsive may suffer in the face of continued climate change. Comparative studies of phenology have found phylogeny to be a reliable predictor of mean leaf‐out date and flowering time at both the local and global scales. This is less true for flowering time response (i.e., the correlation between phenological timing and climate factors), while no study to date has explored whether the response of leaf‐out date to climate factors exhibits phylogenetic signal. We used a 52‐year observational phenological dataset for 52 woody species from the Forest Botanical Garden of Heilongjiang Province, China, to test phylogenetic signal in leaf‐out date and flowering time, as well as, the response of these two phenological traits to both temperature and winter precipitation. Leaf‐out date and flowering time were significantly responsive to temperature for most species, advancing, on average, 3.11 and 2.87 day/°C, respectively. Both leaf‐out and flowering, and their responses to temperature exhibited significant phylogenetic signals. The response of leaf‐out date to precipitation exhibited no phylogenetic signal, while flowering time response to precipitation did. Native species tended to have a weaker flowering response to temperature than non‐native species. Earlier leaf‐out species tended to have a greater response to winter precipitation. This study is the first to assess phylogenetic signal of leaf‐out response to climate change, which suggests, that climate change has the potential to shape the plant communities, not only through flowering sensitivity, but also through leaf‐out sensitivity.     

European phenological response to climate change matches the warming pattern

Global climate change impacts can already be tracked in many physical and biological systems; in particular, terrestrial ecosystems provide a consistent picture of observed changes. One of the preferred indicators is phenology, the science of natural recurring events, as their recorded dates provide a high-temporal resolution of ongoing changes. Thus, numerous analyses have demonstrated an earlier onset of spring events for mid and higher latitudes and a lengthening of the growing season. However, published single-site or single-species studies are particularly open to suspicion of being biased towards predominantly reporting climate change-induced impacts. No comprehensive study or meta-analysis has so far examined the possible lack of evidence for changes or shifts at sites where no temperature change is observed. We used an enormous systematic phenological network data set of more than 125 000 observational series of 542 plant and 19 animal species in 21 European countries (1971–2000). Our results showed that 78% of all leafing, flowering and fruiting records advanced (30% significantly) and only 3% were significantly delayed, whereas the signal of leaf colouring/fall is ambiguous. We conclude that previously published results of phenological changes were not biased by reporting or publication predisposition: the average advance of spring/summer was 2.5 days decade⁻¹ in Europe. Our analysis of 254 mean national time series undoubtedly demonstrates that species' phenology is responsive to temperature of the preceding months (mean advance of spring/summer by 2.5 days°C⁻¹, delay of leaf colouring and fall by 1.0 day°C⁻¹). The pattern of observed change in spring efficiently matches measured national warming across 19 European countries (correlation coefficient r =−0.69, P <0.001).     

Prediction of Arctic plant phenological sensitivity to climate change from historical records

The pace of climate change in the Arctic is dramatic, with temperatures rising at a rate double the global average. The timing of flowering and fruiting (phenology) is often temperature dependent and tends to advance as the climate warms. Herbarium specimens, photographs, and field observations can provide historical phenology records and have been used, on a localised scale, to predict species’ phenological sensitivity to climate change. Conducting similar localised studies in the Canadian Arctic, however, poses a challenge where the collection of herbarium specimens, photographs, and field observations have been temporally and spatially sporadic. We used flowering and seed dispersal times of 23 Arctic species from herbarium specimens, photographs, and field observations collected from across the 2.1 million km² area of Nunavut, Canada, to determine (1) which monthly temperatures influence flowering and seed dispersal times; (2) species’ phenological sensitivity to temperature; and (3) whether flowering or seed dispersal times have advanced over the past 120 years. We tested this at different spatial scales and compared the sensitivity in different regions of Nunavut. Broadly speaking, this research serves as a proof of concept to assess whether phenology–climate change studies using historic data can be conducted at large spatial scales. Flowering times and seed dispersal time were most strongly correlated with June and July temperatures, respectively. Seed dispersal times have advanced at double the rate of flowering times over the past 120 years, reflecting greater late‐summer temperature rises in Nunavut. There is great diversity in the flowering time sensitivity to temperature of Arctic plant species, suggesting climate change implications for Arctic ecological communities, including altered community composition, competition, and pollinator interactions. Intraspecific temperature sensitivity and warming trends varied markedly across Nunavut and could result in greater changes in some parts of Nunavut than in others.     

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.     

Dispersal response to climate change: scaling down to intraspecific variation

Range shift, a widespread response to climate change, will depend on species abilities to withstand warmer climates. However, these abilities may vary within species and such intraspecific variation can strongly impact species responses to climate change. Facing warmer climates, individuals should disperse according to their thermal optimum with consequences for species range shifts. Here, we studied individual dispersal of a reptile in response to climate warming and preferred temperature using a semi‐natural warming experiment. Individuals with low preferred temperatures dispersed more from warmer semi‐natural habitats, whereas individuals with higher preferred temperatures dispersed more from cooler habitats. These dispersal decisions partly matched phenotype‐dependent survival rates in the different thermal habitats, suggesting adaptive dispersal decisions. This process should result into a spatial segregation of thermal phenotypes along species moving ranges which should facilitate local adaptation to warming climates. We therefore call for range shift models including intraspecific variation in thermal phenotype and dispersal decision.     

Latitudinal patterns in the phenological responses of leaf colouring and leaf fall to climate change in Japan

Aim  To estimate the potential effect of global climate change on the phenological responses of plants it is necessary to estimate spatial variations at larger scales. However, previous studies have not estimated latitudinal patterns in the phenological response directly. We hypothesized that the phenological response of plants varies with latitude, and estimated the phenological response to long-term climate change using autumn phenological events that have been delayed by recent climate change.
Location  Japan.
Methods  We used a 53-year data set to document the latitudinal patterns in the climate responses of the timing of autumn leaf colouring and fall for two tree species over a wide range of latitudes in Japan (31 to 44° N). We calculated single regression slopes for leaf phenological timing and air temperature across Japan and tested their latitudinal patterns using regression models. The effects of latitude, time and their interaction on the responses of the phenological timings were also estimated using generalized linear mixed models.
Results  Our results showed that single regression slopes of leaf phenological timing and air temperature in autumn were positive at most stations. Higher temperatures can delay the timing of leaf phenology. Negative relationships were found between the phenological response of leaves to temperature and latitude. Single regression slopes of the phenological responses at lower latitudes were larger than those at higher latitudes.
Main conclusions  We found negative relationships between leaf phenological responsiveness and latitude. These findings will be important for predicting phenological timing with global climate change.     

Evolution in response to climate change: In pursuit of the missing evidence

Climate change is imposing intensified and novel selection pressures on organisms by altering abiotic and biotic environmental conditions on Earth, but studies demonstrating genetic adaptation to climate change mediated selection are still scarce. Evidence is accumulating to indicate that both genetic and ecological constrains may often limit populations' abilities to adapt to large scale effects of climate warming. These constraints may predispose many organisms to respond to climate change with range shifts and phenotypic plasticity, rather than through evolutionary adaptation. In general, broad conclusions about the role of evolutionary adaptation in mitigating climate change induced fitness loss in the wild are as yet difficult to make. Editor's suggested further reading in BioEssays: How will fish that evolved at constant sub‐zero temperatures cope with global warming? Notothenioids as a case study Abstract     

Herbarium records identify sensitivity of flowering phenology of eucalypts to climate: Implications for species response to climate change

Flowering phenology is very sensitive to climate and with increasing global warming the flowering time of plants is shifting to earlier or later dates. Changes in flowering times may affect species reproductive success, associated phenological events, species synchrony, and community composition. Long‐term data on phenological events can provide key insights into the impacts of climate on phenology. For Australia, however, limited data availability restricts our ability to assess the impacts of climate change on plant phenology. To address this limitation other data sources must be explored such as the use of herbarium specimens to conduct studies on flowering phenology. This study uses herbarium specimens for investigating the flowering phenology of five dominant and commercially important Eucalyptus species of south‐eastern Australia and the consequences of climate variability and change on flowering phenology. Relative to precipitation and air humidity, mean temperature of the preceding 3 months was the most influential factor on the flowering time for all species. In response to a temperature increment of 1°C, a shift in the timing of flowering of 14.1–14.9 days was predicted for E. microcarpa and E. tricarpa while delays in flowering of 11.3–15.5 days were found for E. obliqua, E. radiata and E. polyanthemos. Eucalyptus polyanthemos exhibited the greatest sensitivity to climatic variables. The study demonstrates that herbarium data can be used to detect climatic signals on flowering phenology for species with a long flowering duration, such as eucalypts. The robust relationship identified between temperature and flowering phenology indicates that shifts in flowering times will occur under predicted climate change which may affect reproductive success, fitness, plant communities and ecosystems.     

Morphological variation in salamanders and their potential response to climate change

Despite the recognition that some species might quickly adapt to new conditions under climate change, demonstrating and predicting such a fundamental response is challenging. Morphological variations in response to climate may be caused by evolutionary changes or phenotypic plasticity, or both, but teasing apart these processes is difficult. Here, we built on the number of thoracic vertebrae (NTV) in ectothermic vertebrates, a known genetically based feature, to establish a link with body size and evaluate how climate change might affect the future morphological response of this group of species. First, we show that in old‐world salamanders, NTV variation is strongly related to changes in body size. Secondly, using 22 salamander species as a case study, we found support for relationships between the spatial variation in selected bioclimatic variables and NTV for most of species. For 44% of species, precipitation and aridity were the predominant drivers of geographical variation of the NTV. Temperature features were dominant for 31% of species, while for 19% temperature and precipitation played a comparable role. This two‐step analysis demonstrates that ectothermic vertebrates may evolve in response to climate change by modifying the number of thoracic vertebrae. These findings allow to develop scenarios for potential morphological evolution under future climate change and to identify areas and species in which the most marked evolutionary responses are expected. Resistance to climate change estimated from species distribution models was positively related to present‐day species morphological response, suggesting that the ability of morphological evolution may play a role for species’ persistence under climate change. The possibility that present‐day capacity for local adaptation might help the resistance response to climate change can be integrated into analyses of the impact of global changes and should also be considered when planning management actions favouring species persistence.     

Genetic variation and population structure in the endangered greater horseshoe bat Rhinolophus ferrumequinum

Following a dramatic decline last century, the British population of the endangered greater horseshoe bat Rhinolophus ferrumequinum is highly fragmented. To examine the consequences of fragmentation and limited dispersal on patterns of genetic structure and variation, we used microsatellite markers to screen bats from around 50% of the known maternity colonies in Britain, and two areas from continental Europe. Analyses revealed that Welsh and English colonies were genetically isolated. This, and lower variability in Britain than north France, may result from either genetic drift, or the species' colonization history. Gene flow among most neighbouring colonies was not generally restricted, with one exception. These findings have important implications for the ongoing conservation management of this species.     

新疆植被生产力与叶面积指数的变化及其对气候的响应

利用美国探路者卫星遥感资料AVHRR LAI和全球生态模式CASA给出的植被净初级生产力资料(NPP)对新疆地区1982~2000年的植被时空变化进行了定量分析,结果表明新疆地区的LAI和NPP的空间分布严格受水分的制约,与气温呈负相关,表现出干旱内陆地区植被受降水控制的地带特征。相对于20世纪80年代,90年代整个新疆出现了变暖的趋势,降水基本也呈现增加的趋势,在42°N以北地区暖湿转型尤其明显,与这种气候型相对应,植被出现了明显的增加趋势,NPP最大增幅可达45gCm-2a-1。但植被对气温和降水的年际变化响应不一样,降水主要是影响植被峰值的起落,而植被在总体演变趋势上却主要受气温控制,3个分区1984~2000年的气温明显上升,而降水变化趋势不明显,植被受气温控制出现了显著的上升趋势(P<0.01)。     

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.     

黄海和东海小黄鱼遗传多样性的RAPD分析

小黄鱼（Pseudosciaena polyactis）是我国近海重要经济鱼类之一。本文分析了采自黄海和东海5个海区共计48个个体小黄鱼的随机扩增DNA多态性（RAPD）。从40个10 bp引物中选取20个用于群体遗传多样性分析,共检测出145个位点,其中132个（91.03%） 显多态性。用Shannon多样性指数量化的平均遗传多态度为1.93（1.50～2.44）,群体内和群体间的遗传变异比例分别为69%和31%;群体间的平均遗传相似度和遗传距离分别为0.9139和0.0861。用非加权配对算数平均法（UPGMA）聚类分析的结果表明,所分析的5个群体可分为3个地理群系,从分子水平上支持了过去有关学者把黄海和东海的小黄鱼划分为北、中、南3个地理群系的观点。     

Community structure and composition in response to climate change in a temperate steppe

Climate change would have profound influences on community structure and composition, and subsequently has impacts on ecosystem functioning and feedback to climate change. A field experiment with increased temperature and precipitation was conducted to examine effects of experimental warming, increased precipitation and their interactions on community structure and composition in a temperate steppe in northern China since April 2005. Increased precipitation significantly stimulated species richness and coverage of plant community. In contrast, experimental warming markedly reduced species richness of grasses and community coverage. Species richness was positively dependent upon soil moisture (SM) across all treatments and years. Redundancy analysis (RDA) illustrated that SM dominated the response of community composition to climate change at the individual level, suggesting indirect effects of climate change on plant community composition via altering water availability. In addition, species interaction also mediated the responses of functional group coverage to increased precipitation and temperature. Our observations revealed that both abiotic (soil water availability) and biotic (interspecific interactions) factors play important roles in regulating plant community structure and composition in response to climate change in the semiarid steppe. Therefore these factors should be incorporated in model predicting terrestrial vegetation dynamics under climate change.     

Genetic diversity and fitness in Scabiosa columbaria in the Swiss Jura in relation to population size

Due to changes in land use, remnants ofunfertilised, nutrient-poor calcareousgrassland in the Swiss Jura are small in areaand are highly fragmented. We selected 89 seedfamilies from eleven populations of varioussizes of Scabiosa columbaria for a studyof molecular diversity, and used the samematerial in a greenhouse experiment to measurevariation in fitness-related traits and theability of populations to cope withcompetition. Using RAPD-PCR we detected 71RAPD-phenotypes among 87 genotypes. Moleculardiversity within populations was variable andrelatively high, with an expectedheterozygosity H _e ranging from 0.09 to0.24. H _e, the Shannon index (SI) and thepercentage of polymorphic bands were notcorrelated with population size, but thesmallest populations had the lowest moleculardiversity (H _e, SI). Populationdifferentiation was moderate with 12% of themolecular diversity among populations. Measuresof fitness in the greenhouse differed amongseed families (P < 0.001), but not amongpopulations. Mean above-ground biomass waslargely reduced when plants had to compete withBromus. Mean fitness of populationsdecreased when molecular diversity (H _e)was low, but only when plants had to competewith Bromus (P = 0.02). Accordingly, therelative competition ability of Scabiosaplants decreased when molecular diversity(H _e) was low (P = 0.01). Our resultssuggest an increased risk of local extinctionof Scabiosa columbaria in the Swiss Juracaused by a decreased viability and reducedphenotypic plasticity due to genetic erosion insmall populations.     

Genetic response to rapid climate change: it's seasonal timing that matters

The primary nonbiological result of recent rapid climate change is warming winter temperatures, particularly at northern latitudes, leading to longer growing seasons and new seasonal exigencies and opportunities. Biological responses reflect selection due to the earlier arrival of spring, the later arrival of fall, or the increasing length of the growing season. Animals from rotifers to rodents use the high reliability of day length to time the seasonal transitions in their life histories that are crucial to fitness in temperate and polar environments: when to begin developing in the spring, when to reproduce, when to enter dormancy or when to migrate, thereby exploiting favourable temperatures and avoiding unfavourable temperatures. In documented cases of evolutionary (genetic) response to recent, rapid climate change, the role of day length (photoperiodism) ranges from causal to inhibitory; in no case has there been demonstrated a genetic shift in thermal optima or thermal tolerance. More effort should be made to explore the role of photoperiodism in genetic responses to climate change and to rule out the role of photoperiod in the timing of seasonal life histories before thermal adaptation is assumed to be the major evolutionary response to climate change.     

Genetic diversity does not explain variation in extra‐pair paternity in multiple populations of a songbird

Many songbirds are socially monogamous but genetically polyandrous, mating with individuals outside their pair bonds. Extra‐pair paternity (EPP) varies within and across species, but reasons for this variation remain unclear. One possible source of variation is population genetic diversity, which has been shown in interspecific meta‐analyses to correlate with EPP but which has limited support from intraspecific tests. Using eight populations of the genetically polyandrous red‐winged blackbird (Agelaius phoeniceus), including an island population, we investigated whether population‐level differences in genetic diversity led to differences in EPP. We first measured genetic diversity over 10 microsatellite loci and found, as predicted, low genetic diversity in the island population. Additional structure analyses with multilocus genotypes and mtDNA showed the island population to be distinct from the continental populations. However, the island population's EPP rate fell in the middle of the continental populations' distribution, whereas the continental populations themselves showed significant variation in EPP. This result suggests that genetic diversity by itself is not a predictor of EPP rate. We discuss reasons for the departure from previous results, including hypotheses for EPP that do not solely implicate female‐driven behaviour.     

广东省中山市五桂山土沉香遗传多样性

土沉香(Aquilaria sinensis)是生产中药沉香的重要树种,现被列为国家二级重点保护野生植物。该研究通过采集该地区83株土沉香样品,利用限制性酶切位点相关DNA测序(restriction site-associated DNA sequencing,RAD-seq)方法获得的18个微卫星体作为分子标记,计算了五桂山土沉香种群的整体观测杂合度(observed-heterozygosity)、期望杂合度(expected-heterozygosity)、种群整体近交系数及其个体间的亲缘关系,并对种群可能经历的种群缩小或种群扩张状况进行了分析。结果表明:与同属其它物种以及同一生活史物种相比,五桂山地区土沉香遗传多样性略低(观测杂合度和期望杂合度分别为0.523和0.522),但种群整体处于随机交配状态(近交系数为-0.002)。瓶颈效应分析表明该种群没有经历瓶颈效应,且81.810%的个体间无亲缘关系,说明这一地区土沉香并非少数个体发育起来;种群扩张分析表明其种群大小有增长情形。综合这些结果可以看出,中山市五桂山土沉香种群整体遗传健康状况良好,较好地保存了该区域种质资源遗传多样性,为今后的保护和合理开发提供了优质资源储备。     

Genetic variation and population structure of the mosquito Anopheles jeyporiensis in southern China

Genetic differentiation among populations of Anopheles jeyporiensis was examined using 76 mtDNA COII sequences from 16 sites throughout southern China and northern Vietnam. The COII sequences are AT-rich (74.58%) and reveal high levels of diversity with 39 of 685 sites polymorphic and 50 different haplotypes present. Genetic variation is high within populations and significant geographical structure was detected at both population and regional levels. In the larger samples, the distributions of haplotypes suggest recent population expansion.