青藏高原那曲高山嵩草草甸植物物候对增温的异步响应

植物物候对气候变暖的响应是全球气候变化研究的重要内容。目前,高海拔生态系统植物物候对气候变暖响应的研究仍然较少。该研究依托西藏那曲高寒草地生态系统国家野外科学观测研究站布设的梯度增温实验,分别于2015、2017、2018和2021年对模拟增温下优势物种高山嵩草(Kobresia pygmaea)和钉柱委陵菜(Potentilla saundersiana)返青期、现蕾期和开花期等表征植物物候的指标进行了观测,以期揭示增温下藏北高寒草甸植物物候变化机制。结果表明:随着温度升高,高寒草甸中优势植物物候具有不同的变化趋势。高山嵩草返青、现蕾和开花物候期的推迟幅度与温度升高幅度呈正相关关系;钉柱委陵菜返青、现蕾和开花时间随着温度上升表现为先提前后推迟;这表明高寒草甸植物物候对增温产生异步响应。此外,长期增温下的藏北高寒草甸优势种的物候变化均显示出了延迟效应。结构方程归因分析发现,空气温度升高促使高山嵩草返青时间推迟;低水平增温可以促进钉柱委陵菜物候提前,而随着温度继续升高其物候响应发生逆转,土壤水分在决定物候对气候变暖响应的幅度和方向上具有关键作用。该研究结果揭示了藏北高寒草甸优势植物物候响...     

东北大兴安岭演替初期泰加林灌草层典型植物开花物候与生长对模拟暖干化气候的响应

为了揭示暖干化气候变化趋势对演替初期泰加林灌草层植物开花物候和生长的影响, 以东北大兴安岭演替初期泰加林灌草层(高度<50 cm)植物为研究对象, 根据植物生活史, 选择典型的早花植物圆锥薹草(Carex diandra)和水葡萄茶藨子(Ribes procumbens), 中花植物杜香(Ledum palustre)及晚花植物齿叶风毛菊(Saussurea neoserrata), 监测模拟增温和排水处理2年后其完整的开花物候及生长状况, 以期为认识气候变化下泰加林火后植被恢复演替提供依据。结果显示: 1)模拟增温和排水处理后, 早花植物开花温度限制的提早解除使其花期提前, 而水分限制使得晚花植物花期呈延后趋势, 这可能为中花植物提供更大的生态位空隙, 使其开花数量趋向于增加, 花期提前且呈延长趋势; 2)模拟增温和排水处理后, 圆锥薹草、杜香和齿叶风毛菊的盖度和频度均呈增加的趋势, 以杜香最显著, 而水葡萄茶藨子的盖度和频度均下降; 3)模拟增温与排水处理对植物开花物候的影响无显著交互作用, 但土壤水分降低影响部分植物生长对增温的响应。研究结果表明, 植物开花物候对暖干化气候的响应表现出明显的种间差异, 因植物开花功能群而异。早、中花植物花期趋于提前, 晚花期植物花期趋于延后, 一方面可能导致群落生态位变化, 对种间竞争产生潜在影响, 引发群落组成和结构改变; 另一方面可能成为群落调节其整体物候, 以适应气候变化的重要途径。     

Flowering phenology and growth of typical shrub grass plants in response to simulated warmer and drier climate in early succession Taiga forests in the Da Hinggan Ling of northeast China

为了揭示暖干化气候变化趋势对演替初期泰加林灌草层植物开花物候和生长的影响, 以东北大兴安岭演替初期泰加林灌草层(高度<50 cm)植物为研究对象, 根据植物生活史, 选择典型的早花植物圆锥薹草(Carex diandra)和水葡萄茶藨子(Ribes procumbens), 中花植物杜香(Ledum palustre)及晚花植物齿叶风毛菊(Saussurea neoserrata), 监测模拟增温和排水处理2年后其完整的开花物候及生长状况, 以期为认识气候变化下泰加林火后植被恢复演替提供依据。结果显示: 1)模拟增温和排水处理后, 早花植物开花温度限制的提早解除使其花期提前, 而水分限制使得晚花植物花期呈延后趋势, 这可能为中花植物提供更大的生态位空隙, 使其开花数量趋向于增加, 花期提前且呈延长趋势; 2)模拟增温和排水处理后, 圆锥薹草、杜香和齿叶风毛菊的盖度和频度均呈增加的趋势, 以杜香最显著, 而水葡萄茶藨子的盖度和频度均下降; 3)模拟增温与排水处理对植物开花物候的影响无显著交互作用, 但土壤水分降低影响部分植物生长对增温的响应。研究结果表明, 植物开花物候对暖干化气候的响应表现出明显的种间差异, 因植物开花功能群而异。早、中花植物花期趋于提前, 晚花期植物花期趋于延后, 一方面可能导致群落生态位变化, 对种间竞争产生潜在影响, 引发群落组成和结构改变; 另一方面可能成为群落调节其整体物候, 以适应气候变化的重要途径。     

模拟增温改变川西高山草甸优势植物繁殖物候序列特征

以西南横断山区高山草甸优势植物种珠芽拳参(Polygonum viviparum)和银叶委陵菜(Potentilla leuconota)为研究对象, 将其物候分为花芽期、开花期、凋谢期和种子成熟期4个阶段, 每个阶段又分为开始、峰值和结束3个状态。采用开顶式增温箱进行模拟增温, 连续增温4年后, 于增温第5年的2016年生长季跟踪调查了模拟增温对珠芽拳参和银叶委陵菜的繁殖物候序列的影响, 以探讨高山植物群落对气候变化的响应过程。结果显示, 模拟增温后: 1)珠芽拳参各物候阶段的持续时间缩短; 除凋谢阶段起始、结束期延迟外, 其他状态均有不同程度的提前; 各阶段的过渡期有不同程度的缩短, 繁殖周期缩短; 2)银叶委陵菜各物候阶段的持续时间延长; 凋谢期结束前各状态(除开花峰值外)表现为不同程度的提前; 各阶段过渡期对增温的响应不一致, 繁殖周期延长。结果表明: 完整的繁殖物候序列能更准确地反映植物物候对气候变暖的响应; 植物对环境变化的响应和应对策略存在种间差异, 这种差异可能会进一步改变植物群落组成和结构。     

高寒矮生嵩草草甸主要植物物候特征对养分和水分添加的响应

植物物候特征对环境条件的季节和年际变化具有较强的指示作用, 因此研究植物物候特征对环境条件变化的响应, 对理解植物和环境之间的相互作用关系、植物的适应机制和生存策略, 以及应对全球变化都具有重要的意义。该研究基于2009-2011年高寒矮生嵩草(Kobresia humilis)草甸养分水分控制实验的植物物候观测数据资料, 采用巢式方差分析、物候指数和聚类分析方法, 开展了高寒矮生嵩草草甸主要植物物候特征对养分和水分添加的响应研究。结果表明: (1)养分添加处理之间植物返青期和枯黄期均无显著差异, 但养分添加中氮磷处理对主要物种作用较明显, 使莎草科、禾本科、杂类草主要代表植物的返青期和枯黄期推迟。(2)增雪处理效应明显, 主要优势物种无论是何种养分添加, 在增雪处理后均表现出花期物候提前的趋势(p < 0.01), 同时增雪处理使杂类草植物返青期显著提前(p < 0.05)。增水处理对植物的作用效果并不一致, 其中垂穗披碱草(Elymus nutans)和双柱头藨草(Scirpus distigmaticus)的枯黄期显著推迟(p < 0.05), 而杂类草枯黄期提前。(3)养分添加后, 不同物种的物候特征表现出显著差异(p < 0.01), 例如雪白委陵菜(Potentilla nivea)枯黄期显著推迟(p < 0.05), 而双柱头藨草的枯黄期显著提前(p < 0.05), 但物种对养分添加响应的差异以植物类群为单位, 禾本科植物表现为返青期推迟, 而莎草科植物表现为返青期提前。(4)矮生嵩草草甸主要植物营养生长期与果后营养期持续天数之间呈负相关关系, 主要植物物候特征经聚类分析可以分为3个类群, 3个类群经氮磷钾、钾和氮钾三个养分添加处理后植物物候特征变化较大。研究表明, 高寒矮生嵩草草甸植物物候特征在物种水平响应和水分添加后的响应表现出较大差异, 而对养分添加的响应不显著。     

Experimental warming changed plants’ phenological sequences of two dominant species in an alpine meadow,western of Sichuan

以西南横断山区高山草甸优势植物种珠芽拳参(Polygonum viviparum)和银叶委陵菜(Potentilla leuconota)为研究对象, 将其物候分为花芽期、开花期、凋谢期和种子成熟期4个阶段, 每个阶段又分为开始、峰值和结束3个状态。采用开顶式增温箱进行模拟增温, 连续增温4年后, 于增温第5年的2016年生长季跟踪调查了模拟增温对珠芽拳参和银叶委陵菜的繁殖物候序列的影响, 以探讨高山植物群落对气候变化的响应过程。结果显示, 模拟增温后: 1)珠芽拳参各物候阶段的持续时间缩短; 除凋谢阶段起始、结束期延迟外, 其他状态均有不同程度的提前; 各阶段的过渡期有不同程度的缩短, 繁殖周期缩短; 2)银叶委陵菜各物候阶段的持续时间延长; 凋谢期结束前各状态(除开花峰值外)表现为不同程度的提前; 各阶段过渡期对增温的响应不一致, 繁殖周期延长。结果表明: 完整的繁殖物候序列能更准确地反映植物物候对气候变暖的响应; 植物对环境变化的响应和应对策略存在种间差异, 这种差异可能会进一步改变植物群落组成和结构。     

模拟增温改变青藏高原植物繁殖物候及植株高度

气候变化显著影响了高寒植物物候期及生长模式, 从而改变了高寒生态系统功能。而高寒植物物候期和生长状况对气候变化的响应程度, 与其自身资源分配策略有关。为了更好地探究气候变化下高寒植物繁殖物候及生长的规律, 该研究以青藏高原高寒草甸为研究对象, 按生物量从高到低选取15种常见植物, 其生物量之和占样地总生物量80%以上, 采用红外辐射器模拟增温的方法, 利用同质园实验, 观测无种间竞争条件下, 增温2年间植物返青、现蕾、开花以及结实物候, 并监测了植株高度。研究结果表明: (1)在功能群水平上, 增温使豆科类植物的返青、现蕾和开花时间分别显著提前了(8.21 ± 1.81)、(9.14 ± 2.41)和(10.14 ± 2.05) d, 使其开花持续时间显著延长了(6.14 ± 1.52) d, 而增温对其他功能群物候事件无显著影响。增温对高寒植物物候的影响存在种间及年际间差异, 但总体上增温使大多数高寒植物繁殖物候提前并且开花持续时间延长, 将更多的资源更多地分配到繁殖生长上。(2)增温显著降低了杂类草植物的植株高度(平均降低(3.58 ± 0.96) cm), 但对豆科类、禾草类及莎草类功能群植株高度没有显著影响。 增温对高寒植物植株高度的影响存在显著的种间差异以及年际差异。综上所述, 未来气候变暖背景下, 青藏高原高寒植物群落可能更早进入繁殖阶段, 从而降低在营养生长上的资源分配。另外, 由于各物种繁殖能力和营养生长对温度变化响应的差异, 气候变暖将导致高寒植物群落中各物种盖度的变化, 进而改变群落物种组成, 从而影响高寒生态系统的功能。     

川西北高寒草地3种主要植物的生长及物质分配对温度升高的响应

在野外自然条件下采用开顶式生长室模拟增温的方法, 研究了增温对川西北高寒草地3种主要植物(单子叶草本植物垂穗披碱草(Elymus nutans)和双子叶草本植物尼泊尔酸模(Rumex acetosa)和鹅绒委陵菜(Potentilla anserina))的生长及物质分配的影响。研究结果表明, 增温对3种植物的生长产生了显著影响, 垂穗披碱草和尼泊尔酸模的比叶面积和生物量积累在增温后显著增加, 而鹅绒委陵菜在增温后显著减少。在各组分中, 增温处理使尼泊尔酸模的叶生物量显著增加, 根生物量却显著下降, 而鹅绒委陵菜叶和茎的生物量在增温后显著减少, 根生物量却显著增加。增温对尼泊尔酸模各组分的养分含量产生了显著影响, 其中, 根部碳含量在增温后显著增加, 而氮含量在增温后显著减少。增温对尼泊尔酸模和鹅绒委陵菜的生物量在各组分中的分配产生了显著影响, 增温显著增加了尼泊尔酸模的叶重比(LMR)、根重比(RMR)和地下生物量/地上生物量(R/S), 而茎重比(SMR)在增温后却显著降低; 增温显著增加了鹅绒委陵菜的RMR和R/S, 而SMR和LMR在增温后却显著降低。增温对尼泊尔酸模和鹅绒委陵菜各组分中的碳、氮分配产生了显著影响, 增温显著增加了碳、氮在尼泊尔酸模叶片的分配比例, 并且使尼泊尔酸模根部的碳分配比例也显著增加, 而茎部的碳、氮分配比例却显著减少; 增温显著减少了碳在鹅绒委陵菜叶片的分配比例, 而根部的碳、氮分配比例却显著增加。     

Seasonal dynamic of plant phenophases in Beijing—A case study in Beijing Botanical Garden

季节性是北京植物景观的典型特征, 而个体物候是植物景观季节动态的基础。该研究基于北京植物园内120种落叶树种的周年物候数据, 对北京植物景观的物候季节动态进行分析。物候观测以中国物候观测法为标准, 采用a、b、c三级物候代码进行物候记录; 数据分析以二十四节气中的候(5日)为基本时间变量, 基于频率分布型法探究北京物候季节划分及其物候构成动态, 基于SPSS 20.0频数分布统计等探究各类物候现象发生期及持续期的时间分布特征等。物候季节划分及物候构成特征结果为: 6-19候为春, 物候期发生频数占全年总量的54.03%, 以发芽、展叶、开花为主要物候特征, 后期有少数树种结果; 20-45候为夏, 物候量占全年的12.95%, 此期全部观测树种完成展叶, 春花树种进入结果期, 并有较少夏花开放及秋色叶出现; 46-59候为秋, 物候量占全年的27.19%, 以秋色叶及落叶物候为主并伴有较少结果和开花物候发生; 60候至次年春季起始前为冬, 其中60-72候物候量仅占全年的0.6%, 全为落叶物候。各类物候期的时间分布特征如下: 展叶物候期分布于3-23候, 华北珍珠梅(Sorbaria kirilowii)、旱柳(Salix matsudana)等展叶最早, 展叶期平均持续3.27候。秋色物候期分布于40-63候, 49-56候为最佳观赏期, 蒙椴(Tilia mongolica)、山杏(Armeniaca sibirica)等最早显秋色; 秋色期平均持续8.2候, 卫矛(Euonymus alatus)、接骨木(Sambucus williamsii)等秋色期较长。叶幕期平均持续44.2候, 糯米条(Abelia chinensis)、旱柳、棣棠(Kerria japonica)等叶幕期最长。花物候期分布于1-53候, 始花期为1-41候, 盛花期平均发生于始花后1.81候, 8-23候为集中观赏期, 蜡梅(Chimonanthus praecox)、迎春(Jasminum nudiflorum)、榆(Ulmus pumila)、毛白杨(Populus tomentosa)等开花最早, 木香薷(Elsholtzia stauntoni)开花最晚; 花期平均持续 6.7候, 华北珍珠梅、木槿(Hibiscus syriacus)、紫薇(Lagerstroemia indica)等夏秋开花树种花期最长。果物候期分布于8-59候, 榆、郁香忍冬(Lonicera fragrantissima)等果实成熟最早; 持果期平均持续29.0候, 果实宿存树种及黑果荚蒾(Viburnum melanocarpum)、‘金叶’风箱果(Physocarpus opulifolius ‘Luteus’)等具有较长的果实观赏期。与20世纪80年代同类研究结果对比分析, 北京2017年的物候季节与30年前相比, 入春提早1候, 夏季延长4候, 入秋延后3候, 秋季缩短2候, 且不同季节持续期长短的差距加大。     

北京植物物候的季节动态特征——以北京植物园为例

季节性是北京植物景观的典型特征, 而个体物候是植物景观季节动态的基础。该研究基于北京植物园内120种落叶树种的周年物候数据, 对北京植物景观的物候季节动态进行分析。物候观测以中国物候观测法为标准, 采用a、b、c三级物候代码进行物候记录; 数据分析以二十四节气中的候(5日)为基本时间变量, 基于频率分布型法探究北京物候季节划分及其物候构成动态, 基于SPSS 20.0频数分布统计等探究各类物候现象发生期及持续期的时间分布特征等。物候季节划分及物候构成特征结果为: 6-19候为春, 物候期发生频数占全年总量的54.03%, 以发芽、展叶、开花为主要物候特征, 后期有少数树种结果; 20-45候为夏, 物候量占全年的12.95%, 此期全部观测树种完成展叶, 春花树种进入结果期, 并有较少夏花开放及秋色叶出现; 46-59候为秋, 物候量占全年的27.19%, 以秋色叶及落叶物候为主并伴有较少结果和开花物候发生; 60候至次年春季起始前为冬, 其中60-72候物候量仅占全年的0.6%, 全为落叶物候。各类物候期的时间分布特征如下: 展叶物候期分布于3-23候, 华北珍珠梅(Sorbaria kirilowii)、旱柳(Salix matsudana)等展叶最早, 展叶期平均持续3.27候。秋色物候期分布于40-63候, 49-56候为最佳观赏期, 蒙椴(Tilia mongolica)、山杏(Armeniaca sibirica)等最早显秋色; 秋色期平均持续8.2候, 卫矛(Euonymus alatus)、接骨木(Sambucus williamsii)等秋色期较长。叶幕期平均持续44.2候, 糯米条(Abelia chinensis)、旱柳、棣棠(Kerria japonica)等叶幕期最长。花物候期分布于1-53候, 始花期为1-41候, 盛花期平均发生于始花后1.81候, 8-23候为集中观赏期, 蜡梅(Chimonanthus praecox)、迎春(Jasminum nudiflorum)、榆(Ulmus pumila)、毛白杨(Populus tomentosa)等开花最早, 木香薷(Elsholtzia stauntoni)开花最晚; 花期平均持续 6.7候, 华北珍珠梅、木槿(Hibiscus syriacus)、紫薇(Lagerstroemia indica)等夏秋开花树种花期最长。果物候期分布于8-59候, 榆、郁香忍冬(Lonicera fragrantissima)等果实成熟最早; 持果期平均持续29.0候, 果实宿存树种及黑果荚蒾(Viburnum melanocarpum)、‘金叶’风箱果(Physocarpus opulifolius ‘Luteus’)等具有较长的果实观赏期。与20世纪80年代同类研究结果对比分析, 北京2017年的物候季节与30年前相比, 入春提早1候, 夏季延长4候, 入秋延后3候, 秋季缩短2候, 且不同季节持续期长短的差距加大。     

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.     

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.     

Plant functional traits mediate reproductive phenology and success in response to experimental warming and snow addition in Tibet

Global climate change is predicted to have large impacts on the phenology and reproduction of alpine plants, which will have important implications for plant demography and community interactions, trophic dynamics, ecosystem energy balance, and human livelihoods. In this article we report results of a 3‐year, fully factorial experimental study exploring how warming, snow addition, and their combination affect reproductive phenology, effort, and success of four alpine plant species belonging to three different life forms in a semiarid, alpine meadow ecosystem on the central Tibetan Plateau. Our results indicate that warming and snow addition change reproductive phenology and success, but responses are not uniform across species. Moreover, traits associated with resource acquisition, such as rooting depth and life history (early vs. late flowering), mediate plant phenology, and reproductive responses to changing climatic conditions. Specifically, we found that warming delayed the reproductive phenology and decreased number of inflorescences of Kobresia pygmaea C. B. Clarke, a shallow‐rooted, early‐flowering plant, which may be mainly constrained by upper‐soil moisture availability. Because K. pygmaea is the dominant species in the alpine meadow ecosystem, these results may have important implications for ecosystem dynamics and for pastoralists and wildlife in the region.     

Different temperature perception in high‐elevation plants: new insight into phenological development and implications for climate change in the alpine tundra

In alpine habitats, predicted warmer and longer growing seasons will influence plant phenology, with important implications for species adaptation and vegetation dynamics. However, little is known on the temperature sensitivity of different phenophases and on the characteristics allowing phenological variation among and within species. By integrating interannual micro‐climatic variability with experimental warming, we explored how the phenology of three alpine species is influenced by temperature and what mechanisms underlie intra‐ and inter‐specific phenological differences. The present study demonstrated that alpine plants have different temperature responses during their reproductive cycle, do not have constant thermal thresholds and heat‐use efficiencies to achieve the seed dispersal stage and can change their temperature sensitivity to flower along snowmelt gradients. In addition, the length of the reproductive cycle, which proved to be species‐specific under experimental warming, does not seem to be the only life‐history trait under selective pressure due to the short‐length of the snow‐free period. In a warming climate scenario, the phenology of sexual reproduction will be considerably altered, and alpine plants may be subjected to changes in population dynamics driven by altered perception of environmental cues appropriate for coordinating the timing of key life‐history events.     

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.     

Field evidence for earlier leaf-out dates in alpine grassland on the eastern Tibetan Plateau from 1990 to 2006

Worldwide, many plant species are experiencing an earlier onset of spring phenophases due to climate warming. Rapid recent temperature increases on the Tibetan Plateau (TP) have triggered changes in the spring phenology of the local vegetation. However, remote sensing studies of the land surface phenology have reached conflicting interpretations about green-up patterns observed on the TP since the mid-1990s. We investigated this issue using field phenological observations from 1990 to 2006, for 11 dominant plants on the TP at the levels of species, families (Gramineae—grasses and Cyperaceae—sedges) and vegetation communities (alpine meadow and alpine steppe). We found a significant trend of earlier leaf-out dates for one species (Koeleria cristata). The leaf-out dates of both Gramineae and Cyperaceae had advanced (the latter significantly, starting an average of 9 days later per year than the former), but the correlation between them was significant. The leaf-out dates of both vegetation communities also advanced, but the pattern was only significant in the alpine meadow. This study provides the first field evidence of advancement in spring leaf phenology on the TP and suggests that the phenology of the alpine steppe can differ from that of the alpine meadow. These findings will be useful for understanding ecosystem responses to climate change and for grassland management on the TP.     

Experimental warming changed plants' phenological sequences of two dominant species in an alpine meadow,western of Sichuan北大核心CSCD

Aims We studied phenological sequences of two dominant plants (Polygonum viviparum and Potentilla leuconota) in an alpine meadow of the Hengduan Mt., western of Sichuan to explore the alpine plants responses on climate change. Methods Open-Top chambers (OTCs) chosen by ITEX were used to monitor the warming in the field. After a four-year experimental warming, in the 5th growing season we recorded the phenological sequences of two dominant species, focusing on plant responses on warming. The sequence was divided into four stages: budding, flowering, withering and ripe seeds. Each stage had three events: first, peak, and last. Important findings Our results showed that: 1) For P. viviparum, experimental warming elicited a shortening of the duration of each stage, advanced all of the phenological events but the first of withering and ripe seeds, shortened the period of each stage and reduced the duration of entire reproduction. 2) For P. leuconota, experimental warming extended the duration of every stage. All phenological events before the end of withering occurred earlier on experimental warming but the peak of flowering. The period of each stage had inconsistent responses on warming and warming prolonged the duration of entire reproduction. The present results indicated that not all phenological events were equally responsive to experimental warming and an entire sequence could be a more accurate way to evaluate the responses on environmental variation. Therefore, the plastic responses to warming of different species would have effects on community composition and structure. © 2018 Editorial Office of Chinese Journal of Plant Ecology. All rights reserved.     

Flowering phenology and reproductive fitness along a mountain slope: maladaptive responses to transplantation to a warmer climate in Campanula thyrsoides

In many biomes, global warming has resulted in advanced and longer growing seasons, which has often led to earlier flowering in plant taxa. Elevational gradients are ideal to study the effects of global warming as they allow transplantation of plants from their original cooler higher elevations down to elevations with a prospective climate. We transplanted plants from ten populations of the European alpine monocarpic herb species Campanula thyrsoides L. to three sites along a steep mountain slope (600, 1,235 and 1,850 m above sea level) in the Swiss Alps and asked whether reproductive phenology adjusts plastically to elevation and if these responses were adaptive, i.e. increased the fitness of plants. We further assessed current genetic differentiation in phenotypic traits and whether any such origin effects were due to adaptation to climatic conditions of origin. Our results showed that transplantation to lower elevations caused strong shifts in phenology, with plants starting growth and flowering earlier than plants placed at higher elevations. However, compared to flower production at high elevation, number of flowers per plant decreased 21 % at mid- and 61 % at low elevation. The shift in phenology thus came with a high cost in fitness, and we suggest that phenology is maladaptive when C. thyrsoides faces temperature conditions deviating from its natural amplitude. We conclude that the frequently reported phenological shift in plant species as a response to global warming may include heavy fitness costs that may hamper species survival.     

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.