Independent effects of warming and nitrogen addition on plant phenology in the Inner Mongolian steppe

Background and Aims

Phenology is one of most sensitive traits of plants in response to regional climate warming. Better understanding of the interactive effects between warming and other environmental change factors, such as increasing atmosphere nitrogen (N) deposition, is critical for projection of future plant phenology.

Methods

A 4-year field experiment manipulating temperature and N has been conducted in a temperate steppe in northern China. Phenology, including flowering and fruiting date as well as reproductive duration, of eight plant species was monitored and calculated from 2006 to 2009.

Key Results

Across all the species and years, warming significantly advanced flowering and fruiting time by 0·64 and 0·72 d per season, respectively, which were mainly driven by the earliest species (Potentilla acaulis). Although N addition showed no impact on phenological times across the eight species, it significantly delayed flowering time of Heteropappus altaicus and fruiting time of Agropyron cristatum. The responses of flowering and fruiting times to warming or N addition are coupled, leading to no response of reproductive duration to warming or N addition for most species. Warming shortened reproductive duration of Potentilla bifurca but extended that of Allium bidentatum, whereas N addition shortened that of A. bidentatum. No interactive effect between warming and N addition was found on any phenological event. Such additive effects could be ascribed to the species-specific responses of plant phenology to warming and N addition.

Conclusions

The results suggest that the warming response of plant phenology is larger in earlier than later flowering species in temperate grassland systems. The effects of warming and N addition on plant phenology are independent of each other. These findings can help to better understand and predict the response of plant phenology to climate warming concurrent with other global change driving factors.     

内蒙古中西部优势植物春季返青对降水的非线性响应

长期以来,研究植被物候变化与气候因子的关系多基于线性模型,事实上植被物候对气候变化的响应可能是非线性的。以1984-2017年内蒙古中西部温性典型草原和温性草原化荒漠长时间序列植被物候观测资料为基础,分析了近40年来气候变化背景下不同草地类型优势植物返青期变化特征及其对春季降水的非线性响应。结果表明：（1）温性典型草原冷蒿返青期主要受水分控制,与春季降水量表现为开口向下的二次函数关系。气候暖干化导致的水分亏缺是冷蒿返青期呈极显著延迟趋势（1.32 d/a）的主要原因;春季降水量超过60 mm时,冷蒿返青期表现出由延迟转变为提前的趋势。（2）温性草原化荒漠猫头刺返青期主要受热量控制。受春季显著升温影响,猫头刺返青期呈极显著提前趋势（0.63 d/a）。春季降水增多利于猫头刺提早返青,二者表现为开口向上的二次函数关系;春季降水量超过40 mm时,猫头刺返青期对降水的响应程度逐渐降低,这可能与荒漠植物本身的生理特性有关。     

Responses of plant phenology to nitrogen addition: a meta‐analysis

Phenology is one of the most sensitive processes of plant in response to global change. Anthropogenic activities have considerably increased nitrogen (N) deposition, which significant affects plant phenology. Although numerous individual studies have been conducted, it remains controversial how N addition affects phenological stages, and a comprehensive understanding of how plant phenology responds to external N inputs remains elusive. To reconcile the differences, we conducted a meta‐analysis of 117 species to examine the responses of plant phenology to N addition in terrestrial ecosystems, and assessed variations in their responses in relation to ecosystem types, functional groups, and environmental conditions. Our results showed that plant phenology changed significantly after N addition, and phenology time delayed and phenology duration shortened significantly across all biomes except fruiting duration, but varied with biome types. The phenology change in cropland was more dramatical than in grassland after N addition, even in opposite directions. The response of phenological stages to N addition was consistent in two pollination types except the flowering time, the flowering time had no change in anemophilous but significantly delayed in entomophilous. In addition, the response of phenology to N addition was discrepancy among functional groups, the phenology time advanced and duration shortened in sedge, while phenology time delayed and duration shortened in other groups, and the phenology change in legume was larger than grass and forbs. We also found that environmental factors had little effects on the response of plant phenology to N addition, but significant correlation was found between the response ratios of different phenological stages. Our study suggested that phenology was sensitive to N deposition at many phenological stages, and changes in phenology may be smaller with community biodiversity increasing at ecosystem level.     

不同降水条件下内蒙古荒漠草原主要植物物候对长期增温和氮添加的响应

植物物候是生态系统对气候变化响应的重要指示器, 是植物生产力与植被动态模拟的重要参数。但是植物物候对全球变化的响应是否存在年际间变异、年内变异、物种间变异或生境间变异, 以及如何改变, 目前仍然不明确。该研究基于内蒙古荒漠草原长期增温和氮添加实验平台, 选择优势植物短花针茅(Stipa breviflora)、冷蒿(Artemisia frigida)和木地肤(Kochia prostrata)为研究对象, 使用物候打分观测方法和Richards生长曲线拟合方法, 研究了实验处理第11、12和13年(2016-2018年)期间增温和氮添加对植物物候的影响。研究结果表明: (1)短花针茅开花时间集中在第129-145天, 冷蒿开花时间集中在第230-248天, 木地肤开花时间集中在第194-222天。增温、氮添加和增温+氮添加均使短花针茅和木地肤开花时间趋于提前, 冷蒿开花时间趋于推迟。(2)短花针茅结果时间集中在第134-148天, 冷蒿结果时间集中在第241-260天, 木地肤结果时间集中在第207-231天。增温、氮添加和增温+氮添加处理均使短花针茅和木地肤结果时间趋于提前, 冷蒿结果时间趋于推迟。(3)短花针茅生殖生长持续12-25天, 冷蒿生殖生长持续48-55天, 木地肤生殖生长持续45-77天。增温、增温+氮添加缩短了短花针茅而延长了冷蒿和木地肤的生殖生长时间。     

Climate changes affecting biotic interactions,phenology, and reproductive success in a savanna community over a 10-year period

Climatic parameters are able to influence the timing of phenological events affecting the degree of synchrony among plant species, their interactions, and reproductive success. Shrubs of Malpighiaceae family in the Brazilian Tropical Savanna present sequential flowering phenology. We verified variations in climatic factors (temperature and precipitation) over a period of 10 years (2005–2014) and correlated them with the onset of flowering of four of these Malpighiaceae species. Furthermore, we tested whether the phenological synchronization among species has changed over time affecting the herbivory and fruit set. Herbivory and fruit production were recorded during three reproductive seasons (2008/2009, 2011/2012, 2013/2014). We developed a mathematical model to estimate the flower and fruit production in response to phenological changes for the next 5 years. Results show that climatic factors changed, influencing the onset of species flowering. The degree of overlap among species also changed and the effects on species interactions were species specific. The mathematical model successfully presented a tendency on flower and fruit production contributing to the predictions of the outcomes in response to phenological changes. We confirm the effects of climate changes on plant phenological events and the importance of feature plasticity for better performance of species.     

Variable flowering phenology and pollinator use in a community suggest future phenological mismatch

Recent anthropogenic climate change is strongly associated with average shifts toward earlier seasonal timing of activity (phenology) in temperate-zone species. Shifts in phenology have the potential to alter ecological interactions, to the detriment of one or more interacting species. Recent models predict that detrimental phenological mismatch may increasingly occur between plants and their pollinators. One way to test this prediction is to examine data from ecological communities that experience large annual weather fluctuations. Taking this approach, we analyzed interactions over a four-year period among 132 plant species and 665 pollinating insect species within a Mediterranean community. For each plant species we recorded onset and duration of flowering and number of pollinator species. Flowering onset varied among years, and a year of earlier flowering of a species tended to be a year of fewer species pollinating its flowers. This relationship was attributable principally to early-flowering species, suggesting that shifts toward earlier phenology driven by climate change may reduce pollination services due to phenological mismatch. Earlier flowering onset of a species also was associated with prolonged flowering duration, but it is not certain that this will counterbalance any negative effects of lower pollinator species richness on plant reproductive success. Among plants with different life histories, annuals were more severely affected by flowering–pollinator mismatches than perennials. Specialized plant species (those attracting a smaller number of pollinator species) did not experience disproportionate interannual fluctuations in phenology. Thus they do not appear to be faced with disproportionate fluctuations in pollinator species richness, contrary to the expectation that specialists are at greatest risk of losing mutualistic interactions because of climate change.     

Maintenance of temporal synchrony between syrphid flies and floral resources despite differential phenological responses to climate

Variation in species’ responses to abiotic phenological cues under climate change may cause changes in temporal overlap among interacting taxa, with potential demographic consequences. Here, we examine associations between the abiotic environment and plant–pollinator phenological synchrony using a long‐term syrphid fly–flowering phenology dataset (1992–2011). Degree‐days above freezing, precipitation, and timing of snow melt were investigated as predictors of phenology. Syrphids generally emerge after flowering onset and end their activity before the end of flowering. Neither flowering nor syrphid phenology has changed significantly over our 20‐year record, consistent with a lack of directional change in climate variables over the same time frame. Instead we document interannual variability in the abiotic environment and phenology. Timing of snow melt was the best predictor of flowering onset and syrphid emergence. Snow melt and degree‐days were the best predictors of the end of flowering, whereas degree‐days and precipitation best predicted the end of the syrphid period. Flowering advanced at a faster rate than syrphids in response to both advancing snow melt and increasing temperature. Different rates of phenological advancements resulted in more days of temporal overlap between the flower–syrphid community in years of early snow melt because of extended activity periods. Phenological synchrony at the community level is therefore likely to be maintained for some time, even under advancing snow melt conditions that are evident over longer term records at our site. These results show that interacting taxa may respond to different phenological cues and to the same cues at different rates but still maintain phenological synchrony over a range of abiotic conditions. However, our results also indicate that some individual plant species may overlap with the syrphid community for fewer days under continued climate change. This highlights the role of interannual variation in these flower–syrphid interactions and shows that species‐level responses can differ from community‐level responses in nonintuitive ways.     

民勤荒漠植被对气候变化的响应

运用民勤荒漠区1974-2009年物候观测资料和2002-2010年植被样方观测资料以及同期气象资料,分析了荒漠植被对气候变化的响应.结果表明:1961-2010年,民勤荒漠区气温升高,空气湿度增大,年均气温升高速率大于全球水平和中国近百年平均水平;植物对气温变化的响应主要表现在春季物候提前、秋季物候推迟、生长季延长;植被对降水量变化的响应主要表现为植被盖度和纯盖度随降水量减少而降低,植株密度、植物多度随降水量变化而波动;植被盖度和纯盖度与年降水量的相关性较高,然后依次为6-7月和4-5月的降水量;植株密度和植物多度与9月降水量呈正相关;植物春季物候提前的次序是芽初膨大期＞芽开放期＞开花始期＞展叶始期和展叶盛期＞花蕾序出现＞开花盛期＞开花末期＞果实成熟期;秋季物候推迟的次序是叶全变色期＞落叶始期＞叶初变色期＞落叶末期.春季气温升高对民勤荒漠区植物物候的影响大于秋季气温升高对物候的影响.     

中国南亚热带和北温带地区禾本科C3与C4草本植物花果期差异研究

为揭示不同地区禾本科C₃与C₄植物花果期受气候因子的影响,以广东省和内蒙古自治区分别代表南亚热带和北温带地区,从植物志中分别获得两地395和265种禾本科草本植物的3个花果期特征(始花期、末花果期和生殖期长),比较开花物候的差异,并通过一般线性模型探究其与气候因子(年均温与年均降水量)的相关性。结果表明,南亚热带与北温带地区C₃植物的始花期均比C₄植物早。两地C₄共有种在南亚热带地区具有更早的始花期、更晚的末花果期和更长的生殖期,而C₃共有种的末花果期在两地无显著差异,但在南亚热带地区始花期更早,生殖期更长。随年均温升高,北温带地区禾本科植物的始花期提前,而南亚热带地区则延后;随年均降水量升高,两地禾本科植物始花期与末花果期均延迟;禾本科植物生殖期长与年均温和年均降水量均不存在相关性。跨地区分析表明,末花果期、生殖期长与年均温和年均降水量均正相关,而与始花期不相关。禾本科C₃植物比C₄植物对地区间气候差异响应更敏...     

Contrasting effects of warming and increased snowfall on Arctic tundra plant phenology over the past two decades

Recent changes in climate have led to significant shifts in phenology, with many studies demonstrating advanced phenology in response to warming temperatures. The rate of temperature change is especially high in the Arctic, but this is also where we have relatively little data on phenological changes and the processes driving these changes. In order to understand how Arctic plant species are likely to respond to future changes in climate, we monitored flowering phenology in response to both experimental and ambient warming for four widespread species in two habitat types over 21 years. We additionally used long‐term environmental records to disentangle the effects of temperature increase and changes in snowmelt date on phenological patterns. While flowering occurred earlier in response to experimental warming, plants in unmanipulated plots showed no change or a delay in flowering over the 21‐year period, despite more than 1 °C of ambient warming during that time. This counterintuitive result was likely due to significantly delayed snowmelt over the study period (0.05–0.2 days/yr) due to increased winter snowfall. The timing of snowmelt was a strong driver of flowering phenology for all species – especially for early‐flowering species – while spring temperature was significantly related to flowering time only for later‐flowering species. Despite significantly delayed flowering phenology, the timing of seed maturation showed no significant change over time, suggesting that warmer temperatures may promote more rapid seed development. The results of this study highlight the importance of understanding the specific environmental cues that drive species’ phenological responses as well as the complex interactions between temperature and precipitation when forecasting phenology over the coming decades. As demonstrated here, the effects of altered snowmelt patterns can counter the effects of warmer temperatures, even to the point of generating phenological responses opposite to those predicted by warming alone.     

Linking flowering and reproductive allocation in response to nitrogen addition in an alpine meadow

Aims Plants can change in phenology and biomass allocation in response to environmental change. It has been demonstrated that nitrogen is the most limiting resource for plants in many terrestrial ecosystems. Previous studies have usually focused on either flowering phenology or biomass allocation of plants in response to nitrogen addition; however, attempts to link flowering phenology and biomass allocation are still rare. In this study, we tested the effects of nitrogen addition on both flowering phenology and reproductive allocation in 34 common species. We also examined the potential linkage between flowering time and reproductive allocation in response to nitrogen addition.Methods We conducted a 3-year nitrogen addition experiment in Tibetan alpine meadow. We measured first flowering date and the reproductive allocation for 34 common plant species in control, low and high nitrogen added plots, respectively. One-way analysis of variance was used to examine differences of first flowering date and reproductive allocation among treatments. The relationships between the change in species first flowering date and change in reproductive allocation in response to nitrogen addition were examined by calculating Pearson correlation coefficients.Important findings For most species, both first flowering date and reproductive allocation significantly responded to nitrogen addition. Nitrogen addition significantly delayed the first flowering date and reduced the reproductive allocation for all graminoid species, but accelerated flowering and increased reproductive allocation for most forb species. We found that changes in first flowering date significantly negatively correlated with the changes in reproductive allocation over species in response to nitrogen, which indicated a positive relationship between flowering response and plant performance in reproductive allocation. Species that advanced their flowering time with nitrogen addition increased their reproductive allocation, whereas those that delayed flowering time tended to decline in reproductive allocation with nitrogen addition. Our results suggest that species-specific switch from vegetative growth to reproductive growth could influence species performance.     

Phylogenetic conservatism and climate factors shape flowering phenology in alpine meadows

The study of phylogenetic conservatism in alpine plant phenology is critical for predicting climate change impacts; currently we have a poor understanding of how phylogeny and climate factors interactively influence plant phenology. Therefore, we explored the influence of phylogeny and climate factors on flowering phenology in alpine meadows. For two different types of alpine plant communities, we recorded phenological data, including flowering peak, first flower budding, first flowering, first fruiting and the flowering end for 62 species over the course of 5 years (2008–2012). From sequences in two plastid regions, we constructed phylogenetic trees. We used Blomberg’s K and Pagel’s lambda to assess the phylogenetic signal in phenological traits and species’ phenological responses to climate factors. We found a significant phylogenetic signal in the date of all reproductive phenological events and in species’ phenological responses to weekly day length and temperature. The number of species in flower was strongly associated with the weekly day lengths and followed by the weekly temperature prior to phenological activity. Based on phylogenetic eigenvector regression (PVR) analysis, we found a highly shared influence of phylogeny and climate factors on alpine species flowering phenology. Our results suggest the phylogenetic conservatism in both flowering and fruiting phenology may depend on the similarity of responses to external environmental cues among close relatives.     

Effects of experimental warming on plant reproductive phenology in Xizang alpine meadow北大核心CSCD

Aims: Climate warming strongly influences reproductive phenology of plants in alpine and arctic ecosystems. Here we focus on phenological shifts caused by warming in a typical alpine meadow on the Qinghai-Xizang Plateau. Our objective was to explore phenological responses of alpine plant species to experimental warming. Methods: Passive warming was achieved using open-top chambers (OTCs). The treatments included control (C), and four levels of warming (T1, T2, T3, T4). We selected Kobresia pygmaea, Potentilla saundersiana, Potentilla cuneata, Stipa purpurea, Festuca coelestis and Youngia simulatrix as the focal species. Plant phenology was scored every 3-5 days in the growing season. The reproductive phenology phases of each species were estimated through fitting the phenological scores to the Richards function. Important findings: Under soil water stress caused by warming, most plants in the alpine meadow advanced or delayed their reproductive events. As a result, warming significantly delayed phenological development of K. pygmaea. Warming significantly advanced reproductive phenology of P. saundersiana, S. purpurea and F. coelestis, but not of P. cuneata and Y. simulatrix. In addition, warming significantly shortened the average flowering duration of alpine plant species. The potentially warmer and drier growing seasons under climate change may shift the reproductive phenology of the alpine systems in similar pattern.     

Responses of leaf unfolding and flowering to climate change in 12 tropical evergreen broadleaf tree species in Jianfengling,Hainan Island

为了探讨我国热带地区植物物候与气候变化的关系, 利用海南岛尖峰岭热带树木园12种热带常绿阔叶乔木植物2003-2011年物候观测资料结合同期月平均气温和降水数据, 运用积分回归分析方法, 筛选出影响海南岛12种乔木(8种本地种、4种引入种)展叶始期与开花始期的气象因素以及不同气象因素月值变化(月平均气温和月降水量)综合作用对这些树种物候期的动态影响, 最终建立积分回归-物候预测模型, 对气候变化背景下我国热带地区植物物候变化趋势进行了预测。结果表明: 海南岛12种热带常绿阔叶乔木展叶始期与开花始期均对气候变化做出较明显的响应, 几乎所有的树种展叶始期与开花始期的发生都受到气温和降水的共同影响。多数树种展叶始期受展叶前冬季及春季气温影响显著, 且在临近展叶始期的月份, 气温的影响更显著。上一年秋季月降水量对各树种开花始期的影响比其他时段显著, 这验证了降水的滞后性假说。本地种展叶始期对气候变化的响应比其开花始期对气候变化的响应更敏感, 引入种则相反。各树种展叶和开花在受气温和降水综合影响最明显的月份(假设其余11个月份月平均气温和月降水量不变), 月平均气温升高0.1 ℃、月降水量增加10 mm可使展叶始期和开花始期提前或推迟1-3天。积分回归分析方法为解释海南岛热带常绿阔叶乔木物候与气温和降水的动态关系提供了有效的途径, 基于气温和降水与物候资料建立的积分回归-物候预测模型具有对气温和降水变化影响下物候响应的解释率和预测精度高(R²≥ 0.943)的优点, 对于预测气候变化影响下的植物物候变化趋势有一定的适用性。     

海南岛尖峰岭12种热带常绿阔叶乔木展叶期与开花期对气候变化的响应

为了探讨我国热带地区植物物候与气候变化的关系, 利用海南岛尖峰岭热带树木园12种热带常绿阔叶乔木植物2003-2011年物候观测资料结合同期月平均气温和降水数据, 运用积分回归分析方法, 筛选出影响海南岛12种乔木(8种本地种、4种引入种)展叶始期与开花始期的气象因素以及不同气象因素月值变化(月平均气温和月降水量)综合作用对这些树种物候期的动态影响, 最终建立积分回归-物候预测模型, 对气候变化背景下我国热带地区植物物候变化趋势进行了预测。结果表明: 海南岛12种热带常绿阔叶乔木展叶始期与开花始期均对气候变化做出较明显的响应, 几乎所有的树种展叶始期与开花始期的发生都受到气温和降水的共同影响。多数树种展叶始期受展叶前冬季及春季气温影响显著, 且在临近展叶始期的月份, 气温的影响更显著。上一年秋季月降水量对各树种开花始期的影响比其他时段显著, 这验证了降水的滞后性假说。本地种展叶始期对气候变化的响应比其开花始期对气候变化的响应更敏感, 引入种则相反。各树种展叶和开花在受气温和降水综合影响最明显的月份(假设其余11个月份月平均气温和月降水量不变), 月平均气温升高0.1 ℃、月降水量增加10 mm可使展叶始期和开花始期提前或推迟1-3天。积分回归分析方法为解释海南岛热带常绿阔叶乔木物候与气温和降水的动态关系提供了有效的途径, 基于气温和降水与物候资料建立的积分回归-物候预测模型具有对气温和降水变化影响下物候响应的解释率和预测精度高(R²≥ 0.943)的优点, 对于预测气候变化影响下的植物物候变化趋势有一定的适用性。     

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.     

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.     

Effects of experimental warming on plant reproductive phenology in Xizang alpine meadow

Aims Climate warming strongly influences reproductive phenology of plants in alpine and arctic ecosystems. Here we focus on phenological shifts caused by warming in a typical alpine meadow on the Qinghai-Xizang Plateau. Our objective was to explore phenological responses of alpine plant species to experimental warming. Methods Passive warming was achieved using open-top chambers (OTCs). The treatments included control (C), and four levels of warming (T1, T2, T3, T4). We selected Kobresia pygmaea, Potentilla saundersiana, Potentilla cuneata, Stipa purpurea, Festuca coelestis and Youngia simulatrix as the focal species. Plant phenology was scored every 3-5 days in the growing season. The reproductive phenology phases of each species were estimated through fitting the phenological scores to the Richards function. Important findings Under soil water stress caused by warming, most plants in the alpine meadow advanced or delayed their reproductive events. As a result, warming significantly delayed phenological development of K. pygmaea. Warming significantly advanced reproductive phenology of P. saundersiana, S. purpurea and F. coelestis, but not of P. cuneata and Y. simulatrix. In addition, warming significantly shortened the average flowering duration of alpine plant species. The potentially warmer and drier growing seasons under climate change may shift the reproductive phenology of the alpine systems in similar pattern.     

实验增温对藏北高寒草甸植物繁殖物候的影响

全球气候变暖对高寒和极地地区的植物物候产生强烈的影响。该研究主要关注增温条件下藏北高寒草甸不同功能型植物繁殖时间(生殖物候)的改变。实验采用开顶箱式增温方法, 对3个主要功能群浅根-早花、浅根-中花和深根-晚花植物的现蕾、开花、结实时间进行观测。研究结果表明: (1)增温导致了土壤水分胁迫, 显著推迟了浅根-早花植物高山嵩草(Kobresia pygmaea)的繁殖时间; (2)增温显著提前了浅根-中花植物钉柱委陵菜(Potentilla saundersiana)和深根晚花植物紫花针茅(Stipa purpurea)和矮羊茅(Festuca coelestis)的繁殖时间; (3)增温没有显著影响浅根-中花植物楔叶委陵菜(Potentilla cuneata)和深根-晚花植物无茎黄鹌菜(Youngia simulatrix)的繁殖时间; (4)增温缩短了3种类型植物的开花持续时间。这些结果显示增温改变了藏北高寒草甸群落中多数物种的繁殖时间, 这预示着在未来更热更干的生长季, 青藏高原高寒草甸系统的植物物候格局可能会被重塑。     

Later flowering is associated with a compressed flowering season and reduced reproductive output in an early season floral resource

Climate change‐induced shifts in flowering phenology can expose plants to novel biotic and abiotic environments, potentially leading to decreased temporal overlap with pollinators and exposure to conditions that negatively affect fruit and seed set. We explored the relationship between flowering phenology and reproductive output in the common shrub pointleaf manzanita Arctostaphylos pungens in a lower montane habitat in southeastern Arizona, USA. Contrary to the pattern of progressively earlier flowering observed in many species, long‐term records show that A. pungens flowering onset is shifting later and the flowering season is being compressed. This species can thus provide unusual insight into the effects of altered phenology. To determine the consequences of among‐ and within‐plant variation in flowering time, we documented individual flowering schedules and followed the fates of flowers on over 50 plants throughout two seasons (2012 and 2013). We also measured visitation rates by potential pollinators in 2012, as well as both fruit mass and seeds per fruit of flowers produced at different times. Fruit set was positively related to visitation rate but declined with later dates of flower production in both years. Total fruit production per plant was positively influenced by flowering duration, which declined with later flowering onset, as did fruit mass. Individual flowering schedules were consistent between years, suggesting that plants that begin flowering late have lower reproductive output each year. These patterns suggest that if pointleaf manzanita flowering continues to shift later, its flowering season may continue to become shorter, compressing floral resource availability for pollinators and leading to reduced reproductive output. These results reveal the negative effects of delayed phenology on reproductive output in a long‐lived plant. They highlight the value of using natural variation in flowering time, in combination with long‐term data, to anticipate the consequences of phenological shifts.