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地理研究 >

2017 , Vol. 36 >Issue 4: 779 - 789

DOI: https://doi.org/10.11821/dlyj201704015

研究论文

1978-2014年牡丹江地区植物花期变化及模型模拟

  • 徐韵佳 , 1, 2 ,
  • 仲舒颖 1 ,
  • 戴君虎 1 ,
  • 陶泽兴 1, 2 ,
  • 王焕炯 , 1
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  • 1. 中国科学院地理科学与资源研究所,陆地表层格局与模拟重点实验室,北京 100101
  • 2. 中国科学院大学,北京 100049
通讯作者:王焕炯（1987- ）,男,山西榆次人,副研究员,研究方向为物候学与全球变化。E-mail:

作者简介：徐韵佳（1992- ）,女,天津人,硕士,研究方向为气候变化及生态系统响应。E-mail:

收稿日期: 2016-10-21

  要求修回日期: 2017-01-16

  网络出版日期: 2017-04-20

基金资助

国家自然科学基金项目（41401071,41601047）

国家重大科研仪器研制项目（41427805）

中国科学院战略性先导科技专项（XDA05090301）

收起

Changes in flowering phenology of plants and their model simulation in Mudanjiang, China

  • XU Yunjia , 1, 2 ,
  • ZHONG Shuying 1 ,
  • DAI Junhu 1 ,
  • TAO Zexing 1, 2 ,
  • WANG Huanjiong , 1
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  • 1. Key Laboratory of Land Surface Pattern and Simulation, Institute of Geographic Sciences and Natural Resources Research, CAS, Beijing 100101, China
  • 2. University of Chinese Academy of Sciences, Beijing 100049, China

Received date: 2016-10-21

  Request revised date: 2017-01-16

  Online published: 2017-04-20

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摘要

近几十年来,许多研究表明植物始花期随气候变暖普遍提前,但对植物花期长度变化的研究仍较少。利用1978-2014年牡丹江地区40种植物的始花期和末花期观测资料,分析该地区主要植物花期的时间分布、变化特征及与气候变化的关系,并评估两种物候模型对花期模拟的适用性。结果表明：① 牡丹江地区40种植物的花期开始日在4月13日-8月27日之间,结束日在4月25日至9月13日之间,且均集中分布在5月份。花期长度的变化范围在6~69天。大部分植物（62.5%）花期长度在10~20天。② 在研究时段内,植物花期物候出现了一定程度的变化,但大多数植物的变化趋势并不显著。始花期平均推迟速率为0.06天/10a,只有1种植物变化趋势显著（P<0.05）;末花期平均提前速率为0.28天/10a,没有植物变化显著;花期长度平均缩短0.35天/10a,只有4种植物显著缩短。③ 绝大多数植物始花期和末花期的年际变化与季前温度呈显著负相关关系,温度敏感度分别在-6.2天/℃~-2.3天/℃和-5.0天/℃~-1.2天/℃。花期变化趋势不显著与牡丹江地区春季增温趋势不显著有关。④ 回归模型能够准确地模拟始花期、末花期的年际变化,平均拟合优度R2分别为0.65和0.38,对花期长度年际变化的模拟效果稍差（平均R2为0.17）。相比之下,GDD（Growing Degree Days）模型对花期模拟的效果更好,无论是对始、末花期还是花期长度均提高了拟合优度。该研究可为认识植物花期对气候变化的响应以及花期的模拟预报提供依据。

关键词： 牡丹江; 花期长度; 温度敏感度; 物候模型

本文引用格式

徐韵佳 , 仲舒颖 , 戴君虎 , 陶泽兴 , 王焕炯 . 1978-2014年牡丹江地区植物花期变化及模型模拟[J]. 地理研究, 2017 , 36(4) : 779 -789 . DOI: 10.11821/dlyj201704015

Abstract

Over the last several decades, many studies proved that the first flowering date (FFD) of most plant species has become earlier in response to global warming. However, the existing results about the impact of climate change on the flowering duration (FD) were quite limited. In this study, we investigated the temporal distribution and trends in FFD, end of flowering date (EFD) and FD of 40 plants from 1978 to 2014 in Mudanjiang, China. Correlation and regression analyses were used to examine the relationship between the flowering phenophases and the preseason temperature. Meanwhile, we have evaluated the applicability of two phenological models (including the regression model and the growing degree day model) in simulating the flowering phenophases. The results showed that: (1) During the observation period, the mean FFD and EFD of the 40 plant species ranged from April 13 to August 27 and from April 25 to September 13, respectively. However, most of FFD and EFD were concentrated in May. The FD ranged from 6 to 69 days, with most of them (62.5%) ranging between 10 and 20 days. (2) During the study period, FFD became later at a mean rate of 0.06 days decade-1 with only one species showing a significant trend (P<0.05), while EFD occurred earlier at a mean rate of 0.28 days decade-1 (no one significantly). The averaged FD has shortened by 0.35 days decade-1 with only four species showing a significant shortening trend (P<0.05). (3) Most species showed a significant correlation between FFD (or EFD) and preseason temperature. The temperature sensitivities of FFD ranged from -6.2 to -2.3 days/°C-1, while those of EFD varied from -5.0 to -1.2 days/°C-1. The unapparent trends of flowering phenophases were probably due to the unobvious variation of spring temperature in the study area. (4) The regression model has successfully simulated the interannual changes in FFD and EFD with the mean goodness of fit (R2) ranging from 0.38 to 0.65, but failed to simulate FD accurately (mean R2 was 0.17). Compared to the regression model, the growing degree day model has improved the R2 for simulations of FFD, EFD, and FD. Overall, this study provides a basis for a better understanding of flowering phenological responses to climate change and simulation of flowering phenology.

Key words： Mudanjiang; flowering duration; temperature sensitivity; phenological model

1 引言

植物物候变化不仅可以反映全球变化对生物物理系统的影响,还可显著地影响区域和全球尺度的碳循环[1,2]、物种分布范围[3]和动植物种间关系[4,5]。因此,物候学已成为全球变化重点关注的领域之一。花期物候作为植物生殖物候期的一种,主要指植物始花、盛花、末花以及开花持续时间长短等,在植物物候变化研究中占有重要地位[6]。各种不同植物花期相互组合可形成特定的景观,具有独特的美学和经济价值。例如,北京植物园每年举办的桃花节对游客有很强的吸引力,1989-2004年平均每年吸引游客33万人次[7],据国家旅游局统计资料,2014年和2015年接待人数保持在40万以上;樱花节在日本群众的生活中占有举足轻重的地位,日本各地举办樱花节已有千年的历史[8]。另外,因气传致敏花粉的浓度及持续时间与区域的花期物候有关,植物花期变化可能影响到人类健康[9]。因此,研究植物花期物候及对气候变化的响应具有重要的理论意义和应用价值。
国内外大量研究表明,在气候变暖的背景下,北半球春季物候期普遍提前[10-12]。例如,Miller等的研究发现[13],近年来美国东北部地区草本和木本植物的始花期显著提前,温度每升高1 ℃,提前3~8天。在中国,近50年东北和华北地区木本植物的始花期分别以-1.55天/10a和-2.22天/10a的速度提前[14]。近10年来西安地区42种植物的展叶期和始花期相比于1963-1996年分别平均提前了5.54天和10.20天[15]。但现有研究多以不同植物种的始花期为研究对象,较少关注整个花期长度的变化。有个别研究开始注意到花期长度的变化,例如,英国根西岛232种植物的花期长度随气候变暖显著缩短[16],而在其他区域则得到相反的结论。例如,对日本东京97个樱花品种的研究表明大部分植物花期长度呈延长态势[17]。因此,在气候变化背景下花期长度将如何变化还有待深入研究。
选择物候观测资料丰富的牡丹江地区为研究区。首先基于1978-2014年间40种植物的花期观测记录,分析该地区植物始花期、末花期和花期长度的变化趋势。同时,探究花期变化与气候变化间的关系。最后,利用两种通用物候模型模拟这些植物的花期变化,以评估物候模型在花期物候预测中的适用性。

2 研究方法与数据来源

2.1 研究区概况

牡丹江市位于黑龙江省的东南部,总面积约为3.9万km2,境内平均海拔高度230 m。观测点所处的地理位置为44°35′35″N、129°40′31″E（图1）。牡丹江市有松花江最大支流之一的牡丹江流经,地形以中山、低山、丘陵、河谷盆地为主。该地属中温带季风气候,四季分明,夏季暖热多雨,冬季寒冷干燥。1981-2010年的年平均气温为4.8 ℃,全年最冷月是1月,平均气温-16.7 ℃,而最热月为7月,平均气温22.3 ℃（图2）。年平均降水量为561 mm,且年内分布不均匀,主要集中在6-8月,占全年降水量的61%。主要植被类型为柞树林、阔叶混交林及人工落叶松林[18]。
Fig. 1 Geographical location of the study site and its surrounding vegetation

图1 研究区地理位置及周边植被覆盖状况

注：影像获取于2015年5月30日,来源于美国地质调查局。

Fig. 2 Monthly mean temperature and precipitation in Mudanjiang during the period 1981-2010

图2 牡丹江气候概况（以1981-2010年为基准）

2.2 数据来源

所用物候观测资料来源于“中国物候观测网”（China Phenological Observation Network,CPON）牡丹江站点。依据观测年数较长和连续性较好的原则,选择1978-2014年40种植物的始花期（first flowering date,FFD）和末花期（end of flowering date,EFD）序列为研究对象,其中包括38种木本植物,1种藤本植物和1种草本植物（表1）。因部分序列的观测数据不连续,因此观测记录数目在年份间各不相同（图3）,研究时段内平均每年花期记录条数达29条。花期物候的观测均按照中国物候观测网的观测标准[19]来进行。始花期指观测植株上开始出现第一个完全开放花时的日期,而末花期指观测植株上留有极少数花朵时的日期。两者间相差的天数即为花期长度（flowering duration,FD）。
Tab. 1 Summary of the observed species in this study

表1 选择的全部观测植物概况

序号 植物 科 生活型 观测
年数		 平均始
花期		 平均末
花期		 平均花期
长度（天）
1 榛（Corylus heterophylla） 桦木科 落叶灌木或小乔木 25 4-13 4-28 15
2 山杨（Populus davidiana） 杨柳科 落叶乔木 31 4-16 4-29 14
3 榆树（Ulmus pumila） 榆科 落叶乔木 38 4-18 4-25 7
4 春榆（Ulmus davidiana var.japonica) 榆科 落叶乔木 29 4-22 4-29 8
5 落叶松（Larix gmelinii） 松科 落叶乔木 28 4-24 5-4 10
6 梣叶槭（Acer negundo） 槭树科 落叶乔木 30 4-28 5-11 13
7 小青杨（Populus pseudo-simonii） 杨柳科 落叶乔木 20 4-28 5-5 7
8 兴安杜鹃（Rhododendron dauricum） 杜鹃花科 半常绿灌木 29 4-29 5-20 21
9 小叶杨（Populus simonii） 杨柳科 落叶乔木 24 4-30 5-6 6
10 垂柳（Salix babylonica） 杨柳科 落叶乔木 32 5-1 5-15 14
11 山杏（Armeniaca sibirica） 蔷薇科 落叶灌木或小乔木 19 5-1 5-11 10
12 杜梨（Pyrus betulifolia） 蔷薇科 落叶乔木 18 5-3 5-18 15
13 水曲柳（Fraxinus mandschurica） 木犀科 落叶乔木 25 5-3 5-13 9
14 樱桃（Cerasus pseudocerasus） 蔷薇科 落叶乔木 30 5-3 5-18 15
15 李（Prunus salicina） 蔷薇科 落叶乔木 30 5-6 5-19 13
16 黑桦（Betula dahurica） 桦木科 落叶乔木 22 5-7 5-23 16
17 稠李（Padus racemosa） 蔷薇科 落叶乔木 34 5-7 5-21 14
18 朝鲜丁香（Syringa oblata subsp. dilatata） 木犀科 落叶灌木 30 5-8 5-31 23
19 秋子梨（Pyrus ussuriensis） 蔷薇科 落叶乔木 18 5-10 5-24 14
20 欧丁香（Syringa vulgaris） 木犀科 落叶灌木或小乔木 32 5-11 6-2 22
21 山荆子（Malus baccata） 蔷薇科 落叶乔木 35 5-13 5-27 15
22 红皮云杉（Picea koraiensis） 松科 落叶乔木 15 5-14 5-29 15
23 蒙古栎（Quercus mongolica） 壳斗科 落叶乔木 28 5-16 5-22 6
24 花曲柳（Fraxinus rhynchophylla） 木犀科 落叶乔木 15 5-16 5-27 10
25 树锦鸡儿（Caragana arborescens） 豆科 落叶灌木或小乔木 30 5-17 6-7 21
26 卫矛（Euonymus alatus） 卫矛科 落叶灌木 35 5-19 6-6 19
27 樟子松（Pinus sylvestris） 松科 常绿乔木 21 5-20 6-6 16
28 茶条槭（Acer ginnala) 槭树科 落叶灌木或小乔木 33 5-24 6-10 17
29 鼠李（Rhamnus davurica） 鼠李科 落叶灌木或小乔木 29 5-27 6-10 15
30 山里红（Crataegus pinnatifida） 蔷薇科 落叶乔木 36 5-27 6-10 13
31 东北山梅花（Philadelphus schrenkii） 虎耳草科 落叶灌木 29 6-3 6-19 16
32 绿叶悬钩子（Rubus komarovi） 蔷薇科 落叶灌木 26 6-5 6-19 14
33 山葡萄（Vitis amurensis） 葡萄科 落叶藤本 25 6-6 6-18 12
34 花木蓝（Indigofera kirilowii） 豆科 落叶灌木 33 6-11 8-19 69
35 白杜（Euonymus maackii） 卫矛科 落叶乔木 24 6-12 7-3 21
36 刺苞南蛇藤（Celastrus flagellaris） 卫矛科 落叶藤本灌木 23 6-12 6-24 12
37 辽椴（Tilia mandshurica） 椴树科 落叶乔木 27 7-3 7-14 11
38 胡枝子（Lespedeza bicolor） 豆科 落叶灌木 33 7-5 8-31 57
39 朝鲜槐（Maackia amurensis） 豆科 落叶乔木 18 7-6 7-25 19
40 白莲蒿（Artemisia sacrorum） 菊科 落叶半灌木状草本 22 8-27 9-13 16

注：平均始花期和平均末花期以“月-日”的形式表示。

Fig. 3 The number of flowering records for each year in Mudanjiang

图3 牡丹江逐年花期物候记录数量

气象数据下载于中国气象数据网（http://data.cma.cn/）,包括牡丹江1978-2014年日平均气温和降水量资料。通过预分析发现,牡丹江地区四季平均气温在研究时段内呈上升趋势,冬（前一年12月-次年2月）、春季（3-5月）增温趋势不显著;夏（6-8月）、秋季（9-11月）增温显著（P<0.05）,增温速率分别为0.34 ℃/10a和0.45 ℃/10a（图4）。
Fig. 4 The seasonal mean temperature in Mudanjiang from 1978 to 2014

图4 1978-2014年牡丹江四季温度变化

2.3 统计分析与模型模拟

首先,计算各植物物候期（始花期、末花期和花期长度）与年份之间线性回归的斜率来表征1978-2014年间不同物候期的变化趋势,同时分春季开花（始花期在3-5月）和夏季开花（始花期在6-8月）植物进行对比。其次,利用始花期（或末花期）与气温间的相关与回归分析来研究花期变化与气温间的关系。据以往研究[20,21],植物始花期往往与开花前一段时间内的平均气温显著相关,这一时段又被称为最优区间。对每种植物,以多年平均始花期为终点,每隔1天作为一个时段（即始花期前1天,始花期前2天,直至180天）,计算这一时段平均气温与始花期的相关系数,以相关系数绝对值最大的时段作为最优区间。始花期与最优区间内平均气温的回归方程斜率,可表征始花期对温度变化响应的敏感度。对于各植物的末花期而言,最优区间的起点与始花期相同,结束于多年末花期的平均日期。
为研究物候模型在花期预测中的适用性,选取了两种物候模型来模拟40种植物的花期物候。第一种模型为统计模型,即采用一元回归方程分别模拟始花期和末花期,其公式为：
t F = a 1 x 1 + b 1 （1）
t E = a 2 x 2 + b 2 （2）
式中：tF为始花期模拟值（日序）;tE为末花期模拟值（日序）;x1为始花期最优区间的平均气温;x2为末花期最优区间的平均气温;a1、a2为回归系数;b1、b2为截距。
第二种模型为生长度日（growing degree days,GDD）模型。该模型公式为：
∑ t 0 t F R ( x t ) = F FFD （3）
∑ t 0 t E R ( x t ) = F EFD （4）
R ( x t ) = 0 x t ≤ T b x t - T b x t > T b （5）
式中：xt为日平均温度;tF、tE分别为始花期和末花期;t0为有效积温累积的开始日期;Tb为发育基点温度;R(xt)为日积温单元;FFFD和FEFD分别为实现始花期和末花期对应的积温阈值。当积温累积到这一阈值（FFFD、FEFD）时,物候期（tF、tE）来临。该模型需要拟合的参数为t0、Tb、FFFD、FEFD。
对每种植物的始花期或末花期序列,利用所有年的观测数据和日平均气温对两种模型进行参数拟合。参数拟合采用最小二乘法原则,即找到使观测值与模拟值的误差平方和最小的参数组合。花期长度的模拟则通过末花期和始花期模拟值的差值来实现。为检验模型的准确性,分别计算各模型的拟合优度R2和均方根误差（root mean square error,RMSE）。

3 结果分析

3.1 牡丹江地区花期及变化特征

牡丹江地区40种植物的平均始花期在4月13日（榛）至8月27日（白莲蒿）,且有超过一半植物集中在5月（图5a）。平均末花期在4月25日（白榆）至9月13日（白莲蒿）,峰值同样出现在5月（47.5%）。花期长度在6~69天（图5b）,最短的植物为蒙古栎,最长的为花木蓝。大多数植物（62.5%）的花期长度为10~20天。
Fig. 5 Number of species in first flowering date, end of flowering date by month and frequency distribution of flowering duration in Mudanjiang averaged from 1978 to 2014

图5 1978-2014年牡丹江地区40种植物平均始花期、末花期和花期长度的频率分布

1978-2014年,牡丹江地区40种植物始花期发生了一定程度的变化,其变化趋势在-2.34天/10a~2.31天/10a（图6a）。其中,22种植物的始花期提前（55%）,平均提前速率为-0.63天/10a,但趋势显著的只有胡枝子;18种植物的始花期推迟（45%）,平均推迟速率为0.92天/10a,无植物显著。末花期的变化趋势在-3.17天/10a~2.29天/10a之间（图6b）。其中,24种植物的末花期提前（60%）,平均提前速率为-1.14天/10a,其余植物的末花期以平均为1.01天/10a的速度推迟。没有任何一种植物的末花期变化趋势达到了P<0.05的显著度水平。对花期长度而言,25种植物的花期缩短,其中4种植物显著缩短（黑桦、鼠李、绿叶悬钩子和白杜）,且没有植物显著延长（图6c）。对所有植物平均而言,始花期平均推迟0.06天/10a,末花期平均提前0.28天/10a,花期长度平均缩短0.35天/10a。
Fig. 6 Frequency distributions of trends in flowering phenophases of 40 plant species in Mudanjiang from 1978 to 2014

图6 1978-2014年牡丹江地区40种植物花期变化趋势的频率分布

注：虚线表示平均值

春季开花植物与夏季开花植物的花期变化呈不同特征。对于春季开花的30种植物,其平均始花期呈推迟趋势,推迟速率为0.30天/10a,而末花期变化幅度很小,推迟速率仅为0.01天/10a（图6d、图6e）。花期长度平均缩短0.28天/10a（图6f）。与春季开花植物相反,夏季开花植物平均始花期和末花期呈提前趋势,提前速率分别为0.63天/10a和1.18天/10a（图6g、图6h）。夏季开花植物花期长度总体也呈缩短趋势,但缩短速率更快,达0.56天/10a（图6i）。

3.2 气温对花期变化的影响

影响始花期的最优区间长度为30~90天（平均为48天）。绝大多数植物（92.5%）始花期与季前温度（即最优区间内气温）呈显著负相关（P<0.05）,即气温升高会导致始花期提前。其中,白杜的始花期温度敏感度最强（-6.2天/℃）,辽椴最弱（-2.3天/℃）。所有植物始花期温度敏感度的均值为-3.6天/℃。影响末花期的最优区间长度为36~111天（平均值为57天）。77.5%的植物末花期与其季前温度呈显著负相关关系（图7d）。在这些植物中,末花期的温度敏感度变化范围是-5.0（绿叶悬钩子）到-1.2天/℃（欧丁香）。所有植物末花期温度敏感度的均值为-3.3天/℃。
Fig. 7 Frequency distributions of regression slopes of first flowering date and end of flowering date on preseason temperature in Mudanjiang

图7 牡丹江地区40种植物及不同季节始花植物的始花期、末花期与季前气温的回归系数频率分布

注：虚线代表平均值。

不同季节始花的植物对温度变化的响应存在一定差异。春季开花的30种植物始花期平均敏感度为-3.4天/℃,只有两种植物（朝鲜丁香和花曲柳）的始花期与温度不显著相关（图7b）。春季开花植物末花期平均敏感度为-3.3天/℃,有5种植物（朝鲜丁香、花曲柳、樟子松、红皮云杉和树锦鸡儿）的末花期与温度不显著相关（图7e）。夏季开花的10种植物始花期的平均敏感度稍强,达-4.3天/℃,仅有1种植物（白莲蒿）始花期受温度变化影响不显著（图7c）。夏季开花植物的末花期平均敏感度为-3.5天/℃,有4种植物（白莲蒿、花木蓝、胡枝子和朝鲜槐）末花期与温度的相关关系不显著（图7f）。

3.3 花期模拟

针对各植物始花期和末花期建立的回归模型平均能够分别解释65%和38%的始花期和末花期变化（表2）。回归模型对始花期变化的模拟效果较好,平均RMSE为3.34天,而对末花期模拟的RMSE稍大,达5.44天。回归模型对花期长度变化的模拟效果稍差,平均只能解释17%的花期长度年际变化,RMSE为5.05天。
Tab. 2 Validity of regression and growing degree days models for simulating the first floweringdate, end of flowering date and flowering duration

表2 两种模型模拟始花期、末花期和花期长度的检验结果

模型类型 始花期 末花期 花期长度
R2 Sig（%） RMSE R2 Sig（%） RMSE R2 Sig（%） RMSE
回归模型 0.65 93 3.34 0.38 78 5.44 0.17 50 5.05
GDD模型 0.71 93 3.24 0.49 80 5.38 0.27 63 4.62

注：R2代表40个物种平均的拟合优度;Sig代表模拟值与预测值相关性达到显著（P<0.05）的物种占比;RMSE代表平均均方根误差（天）。

采用GDD模型提高了对花期模拟的准确性（表2）。始花期的平均拟合优度从0.65增至0.71,平均RMSE从3.34天下降至3.24天。GDD模型对末花期模拟的提升效果更为明显,平均拟合优度从0.38增至0.49。对花期长度,GDD模型同样显著提升了拟合优度,平均达27%,且模拟值与预测值相关关系显著（P<0.05）的物种百分比从回归模型的50%增至63%。

4 讨论

研究结果表明,牡丹江地区的植物花期发生了一定程度的变化,始花期平均推迟0.06天/10a,末花期平均提前0.28天/10a。其中只有1种植物始花期显著提前,没有植物末花期呈显著变化。绝大部分植物的始花期、末花期与季前温度呈显著负相关关系。本文发现影响花期的春季气温变化趋势不显著（图4b）,这可能是导致花期物候变化趋势不显著的原因。
不同植物种对温度变化的响应程度不同（图7）,这反映了植物对气候变化的适应能力不同[22],同时也体现了不同植物对传粉者等资源的竞争关系在气候变化背景下可能发生改变[23]。尽管不同植物种间始花期、末花期的温度敏感度差异较大,但同一物种始花期和末花期对温度响应的敏感程度差异较小（图8）。大多数植物（67.5%）始花期与末花期温度敏感度差异在±1天/℃之间,平均只有0.18天/℃。这使得在物种水平上仅有4个物种的花期长度显著缩短,总体上花期长度变化并不显著。但花期长度的微弱变化可能造成群落水平上更为显著的变化[24],从而改变植物的授粉成功率,进而影响生态系统结构和功能。
Fig. 8 Frequency distribution of the differences in the temperature sensitivity of first flowering date and end of flowering date for 40 plant species in Mudanjiang

图8 牡丹江地区40种植物的始花期和末花期温度敏感度差值频率分布

注：虚线代表平均值;P&lt;0.05指始、末花期敏感度均显著。

在中国其他地区,植物花期物候也发生了一定变化。例如,北京48种木本植物的始花期在1990-2007年比1963-1989年平均提前了5.4天[25]。Ge等的研究表明中国东部22个站点23种植物的始花期在1963-2006年间以1.21天/10a的速率平均提前了5.2天[26]。这说明,中国东部总体的始花期变化趋势比牡丹江地区更加明显。在中国其他地区,仅有一项研究涉及植物花期长度变化。在西双版纳,雨季日照时长的减少导致5种植物花期长度在1973-1999年间以21天/10a的速率缩短[27],远超过牡丹江地区植物花期长度的变化速度。这表明,花期长度变化存在显著的站点差异。本文中植物始花期与末花期的温度敏感度变化范围分别为-6.2天/℃~-2.3天/℃和-5.0天/℃~-1.2天/℃,这也与其他地区的研究结果一致。例如,Dai等研究表明西安地区42种木本植物始花期受其前期13~194天的温度影响较强,敏感度变化范围为-11.44天/℃~-2.84天/℃[15]。
本文以气温为输入参数的物候模型对植物始花期和末花期的模拟效果良好,表明始花期和末花期的年际变化主要受温度控制。但回归模型对花期长度年际变化的模拟效果较差,这主要是因为在模拟花期长度过程中叠加了始花期和末花期两者的误差。另一方面,引起花期长度变化的驱动因子较为复杂[16]。例如,在热带雨林地区,植物花期长度在干湿季表现出不同的变化特征[28],这表明降水也对花期长度有影响。李军等对中国江苏省苏州市银桂（Osmanthus fragranscv）花期的研究结果表明,银桂初花期与相对湿度关系密切[29]。Pau等表明在植物生长季较长、年际变化较小的地区,物候变化主要由生物因素驱动而不是环境因素[30]。因此,在未来探究花期长度变化机制时,需综合考虑除温度外的降水、湿度、日照时数等环境因素以及物种间相互作用的生物因素[4]。

5 结论

本文揭示了1978-2014年牡丹江地区40种植物花期的时间分布、变化特征及与气候变化的关系,并评估了两种物候模型对花期模拟的适用性,主要得到以下结论：
（1）牡丹江地区40种植物的花期开始于4月13日-8月27日,结束于4月25日-9月13日,始花期和末花期均集中在5月份。平均花期长度在6~69天,但大部分植物花期长度为10~20天。
（2）植物花期出现了一定程度的变化,但大多数植物的变化趋势并不显著。始花期平均推迟0.06天/10a,只有1种植物变化趋势显著;末花期平均提前速率为-0.28天/10a,没有任何一种植物变化显著;花期长度平均缩短0.35天/10a,只有4种植物显著缩短。
（3）绝大多数的植物始花期和末花期的年际变化与温度呈显著负相关关系,温度敏感度分别为-6.2天/℃~-2.3天/℃和-5.0天/℃~-1.2天/℃。花期变化趋势不显著与牡丹江地区春季增温趋势不显著有关。
（4）回归模型能够准确地模拟始花期、末花期的年际变化,但对花期长度年际变化的模拟效果稍差。GDD模型能显著的提升花期模拟效果。因此,在观赏植物花期预报、致敏花粉预报等应用领域,推荐采用GDD模型对花期进行模拟。

The authors have declared that no competing interests exist.

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[10]
Menzel A, Sparks T H, Estrella N, et al.European phenological response to climate change matches the warming pattern. Global Change Biology, 2006, 12(10): 1969-1976.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 decade63641; 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°C63641; delay of leaf colouring and fall by 1.0 day°C63641;). The pattern of observed change in spring efficiently matches measured national warming across 19 European countries (correlation coefficient r=-0.69, P<0.001).

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[11]
Richardson A D, Keenan T F, Migliavacca M, et al.Climate change, phenology, and phenological control of vegetation feedbacks to the climate system. Agricultural and Forest Meteorology, 2013, 169: 156-173.Vegetation phenology is highly sensitive to climate change. Phenology also controls many feedbacks of vegetation to the climate system by influencing the seasonality of albedo, surface roughness length, canopy conductance, and fluxes of water, energy, CO 2 and biogenic volatile organic compounds. In this review, we first discuss the environmental drivers of phenology, and the impacts of climate change on phenology, in different biomes. We then examine the vegetation-climate feedbacks that are mediated by phenology, and assess the potential impact on these feedbacks of shifts in phenology driven by climate change. We finish with an overview of phenological modeling and we suggest ways in which models might be improved using existing data sets. Several key weaknesses in our current understanding emerge from this analysis. First, we need a better understanding of the drivers of phenology, particularly in under-studied biomes (e.g. tropical forests). We do not have a mechanistic understanding of the role of photoperiod, even in well-studied biomes. In all biomes, the factors controlling senescence and dormancy are not well-documented. Second, for the most part (i.e. with the exception of phenology impacts on CO 2 exchange) we have only a qualitative understanding of the feedbacks between vegetation and climate that are mediated by phenology. We need to quantify the magnitude of these feedbacks, and ensure that they are accurately reproduced by models. Third, we need to work towards a new understanding of phenological processes that enables progress beyond the modeling paradigms currently in use. Accurate representation of phenological processes in models that couple the land surface to the climate system is particularly important, especially when such models are being used to predict future climate.

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[12]
Ge Q S, Wang H J, Rutishauser T, et al.Phenological response to climate change in China: A meta-analysis. Global Change Biology, 2015, 21(1): 265-274.Abstract The change in the phenology of plants or animals reflects the response of living systems to climate change. Numerous studies have reported a consistent earlier spring phenophases in many parts of middle and high latitudes reflecting increasing temperatures with the exception of China. A systematic analysis of Chinese phenological response could complement the assessment of climate change impact for the whole Northern Hemisphere. Here we analyse 1263 phenological time series (1960-2011, with 20+ years data) of 112 species extracted from 48 studies across 145 sites in China. Taxonomic groups include trees, shrubs, herbs, birds, amphibians and insects. Results demonstrate that 90.8% of the spring/summer phenophases time series show earlier trends and 69.0% of the autumn phenophases records show later trends. For spring/summer phenophases, the mean advance across all the taxonomic groups was 2.75 days decade(-1) ranging between 2.11 and 6.11 days decade(-1) for insects and amphibians, repsectively. Herbs and amphibians show significantly stronger advancement than trees, shrubs and insect. The response of phenophases of different taxonomic groups in autumn is more complex: trees, shrubs, insects, herbs and insects show a delay between 1.93 and 4.84 days decade(-1) , while other groups reveal an advancement ranging from 1.10 to 2.11 days decade(-1) . For woody plants (including trees and shrubs), the stronger shifts towards earlier spring/summer were detected from the data series starting from more recent decades (1980s-2000s). The geographic factors (latitude, longitude and altitude) could only explain 9% and 3% of the overall variance in spring/summer and autumn phenological trends, respectively. The rate of change in spring/summer phenophase of woody plants (1960s-2000s) generally matches measured local warming across 49 sites in China (R=-0.33, P<0.05). This article is protected by copyright. All rights reserved.

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[13]
Miller-Rushing A J, Primack R B. Global warming and flowering times in Thoreau's Concord: A community perspective. Ecology, 2008, 89(2): 332-341.As a result of climate change, many plants are now flowering measurably earlier than they did in the past. However, some species' flowering times have changed much more than others. Data at the community level can clarify the variation in flowering responses to climate change. In order to determine how North American species' flowering times respond to climate, we analyzed a series of previously unstudied records of the dates of first flowering for over 500 plant taxa in Concord, Massachusetts, USA. These records began with six years of observations by the famous naturalist Henry David Thoreau from 1852 to 1858, continued with 16 years of observations by the botanist Alfred Hosmer in 1878 and 1888-1902, and concluded with our own observations in 2004, 2005, and 2006. From 1852 through 2006, Concord warmed by 2.4 degrees C due to global climate change and urbanization. Using a subset of 43 common species, we determined that plants are now flowering seven days earlier on average than they did in Thoreau's times. Plant flowering times were most correlated with mean temperatures in the one or two months just before flowering and were also correlated with January temperatures. Summer-flowering species showed more interannual variation in flowering time than did spring-flowering species, but the flowering times of spring-flowering species correlated more strongly to mean monthly temperatures. In many cases, such as within the genera Betula and Solidago, closely related, co-occurring species responded to climate very differently from one another. The differences in flowering responses to warming could affect relationships in plant communities as warming continues. Common St. John's wort (Hypericum perforatum) and highbush blueberry (Vaccinium corymbosum) are particularly responsive to changes in climate, are common across much of the United States, and could serve as indicators of biological responses to climate change. We discuss the need for researchers to be aware, when using data sets involving multiple observers, of how varying methodologies, sample sizes, and sampling intensities affect the results. Finally, we emphasize the importance of using historical observations, like those of Thoreau and Hosmer, as sources of long-term data and to increase public awareness of biological responses to climate change.

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[14]
Dai J H, Wang H J, Ge Q S.The decreasing spring frost risks during the flowering period for woody plants in temperate area of eastern China over past 50 years. Journal of Geographical Sciences, 2013, 23(4): 641-652.The temperate monsoon area of China is an important agricultural region but late spring frosts have frequently caused significant damage to plants there. Based on phenological data derived from the Chinese Phenological Observation Network (CPON), corresponding meteorological data from 12 study sites and phenological modeling, changes in flowering times of multiple woody plants and the frequency of frost occurrence were analyzed. Through these analyses, frost risk during the flowering period at each site was estimated. Results of these estimates suggested that first flowering dates (FFD) in the study area advanced significantly from 1963 to 2009 at an average rate of 611.52 days/decade in Northeast China ( P <0.01) and 612.22 days/decade ( P <0.01) in North China. Over the same period, the number of frost days in spring decreased and the last frost days advanced across the study area. Considering both flowering phenology and occurrence of frost, the frost risk index, which measures the percentage of species exposed to frost during the flowering period in spring, exhibited a decreasing trend of 610.37% per decade (insignificant) in Northeast China and 611.80% per decade ( P <0.01) in North China, implying that frost risk has reduced over the past half century. These conclusions provide important information to agriculture and forest managers in devising frost protection schemes in the region.

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[15]
Dai J H, Wang H J, Ge Q S.Multiple phenological responses to climate change among 42 plant species in Xi'an, China. International Journal of Biometeorology, 2013, 57(5): 749-758.Phenology; Climate change; First leaf date; Leaf coloring date; First flowering date; Growing season length

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[16]
Bock A, Sparks T H, Estrella N, et al.Changes in first flowering dates and flowering duration of 232 plant species on the island of Guernsey. Global Change Biology, 2014, 20(11): 3508-3519.Abstract Climate change has affected plant phenology; increasing temperatures are associated with advancing first flowering dates. The impact on flowering duration, however, has rarely been studied. In this study, we analysed first flowering dates and flowering durations from a 27 year dataset of weekly flower observations on 232 plant species from the island of Guernsey in the English Channel. The aim of this study was to explore variation in trends and relationships between first flowering dates, flowering duration and temperature. We specifically looked for evidence that traits, such as life forms and phylogenetic groups, explained variation in sensitivity of first flowering and flowering duration among species. Overall trends revealed significantly earlier flowering over time, by an average of 5.2 days decade(-1) since 1985. A highly significant shortening of flowering duration was observed by an average of 10 days decade(-1) . Correlations between first flowering, flowering duration and year varied between different species, traits and flowering periods. Significant differences among traits were observed for first flowering and to a lesser degree for flowering duration. Overall, in comparison to first flowering, more species had significant trends in flowering duration. Temperature relationships revealed large differences in strength and direction of response. 55% of the species revealed a significant negative relationship of first flowering dates and temperature. In contrast, only 19% of flowering durations had a significant negative temperature relationship. The advance in first flowering date together with a shortening of flowering duration suggests potentially serious impacts on pollinators, which might pose a major threat to biodiversity, agriculture and horticulture. Human health, in terms of pollen allergies, however, might benefit from a shortening of specific plant pollen seasons. 2014 John Wiley & Sons Ltd.

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[17]
Miller-Rushing A J, Katsuki T, Primack R B, et al. Impact of global warming on a group of related species and their hybrids: Cherry tree (Rosaceae) flowering at Mt. Takao, Japan. American Journal of Botany, 2007, 94(9): 1470-1478.Climate change is affecting plant phenology worldwide. Phenological responses vary among species, but it is not clear how responses differ among closely related species. We examined a 25-yr record (1981-2005) of flowering times for 97 trees, representing 17 species and hybrids of cherry (Cerasus sp. or Prunus sp.) grown at Mt. Takao, in Tokyo, Japan. The cherry trees flowered earlier over time, by an average of 5.5 d over the 25-yr study. Earlier flowering was explained largely by a 1.8掳C increase in February-March mean monthly temperatures. Most species and hybrids flowered 3-5 d earlier for each 1掳C increase in temperature, but early-flowering taxa flowered as much as 9 d earlier for each 1掳C increase in temperature. Flowering durations and differences in flowering times among species were greater in warm years than in cold years. Species and individual trees also flowered longer in warm years. These results show that the flowering times of closely related species may change similarly in response to climate change, but that early-flowering species may diverge from the overall trend in a predictable way. Such changes in flowering may affect gene flow and pollination as the length of the flowering season increases.

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[18]
孙元发, 张宇. 牡丹江市乔木植物多样性的研究. 防护林科技, 2013, (6): 69-70.为了准确掌握牡丹江市木本植物多样性的情况,于2010年在牡丹江全市范围内进行了乔木植物多样性的调查。经统计,得到了各物种的分布情况,并通过simpson指数的计算得出牡丹江市乔木植物多样性程度较高的结论。

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[Sun Yuanfa, Zhang Yu.Study of woody plant diversity in Mudanjiang city. Protection Forest Science and Technology, 2013, (6): 69-70.]

[19]
宛敏渭, 刘秀珍. 中国物候观测方法. 北京: 科学出版社, 1979.

[Wan Minwei, Liu Xiuzhen.China's National Phenological Observational Criterion. Beijing: Science Press, 1979.]

[20]
Matsumoto K, Ohta T, Irasawa M, et al.Climate change and extension of the Ginkgo biloba L. growing season in Japan. Global Change Biology, 2003, 9(11): 1634-1642.Abstract To understand the effects of climate change on the growing season of plants in Japan, we conducted trend analysis of phenological phases and examined the relationship between phenology and air temperatures. We used phenological data for Ginkgo biloba L., collected from 1953 to 2000. We defined the beginning and the end of the growing season (BGS and EGS) as the dates of budding and leaf fall, respectively. Changes in the air temperature in the 45 days before the date of BGS affected annual variation in BGS. The annual variation in air temperature over the 85 days before EGS affected the date of EGS. The average annual air temperature in Japan has increased by 1.3°C over the last four decades (1961–2000), and this increase has caused changes in ginkgo phenology. In the last five decades (1953–2000), BGS has occurred approximately 4 days earlier than previously, and EGS has occurred about 8 days later. Consequently, since 1953 the length of the growing season (LGS) has been extended by 12 days. Since around 1970, LGS and air temperatures have shown increasing trends. Although many researchers have stated that phenological events are not affected by the air temperature in the fall, we found high correlations not only between budding dates and air temperatures in spring but also between leaf-fall dates and air temperatures in autumn. If the mean annual air temperature increases by 1°C, LGS could be extended by 10 days. We also examined the spatial distribution of the rate of LGS extension, but we did not find an obvious relationship between LGS extension and latitude.

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[21]
Dai J H, Wang H J, Ge Q S.The spatial pattern of leaf phenology and its response to climate change in China. International Journal of Biometeorology, 2014, 58(4): 521-528.Leaf phenology has been shown to be one of the most important indicators of the effects of climate change on biological systems. Few such studies have, however, been published detailing the relationship between phenology and climate change in Asian contexts. With the aim of quantifying species' phenological responsiveness to temperature and deepening understandings of spatial patterns of phenological and climate change in China, this study analyzes the first leaf date (FLD) and the leaf coloring date (LCD) from datasets of four woody plant species, Robinia pseudoacacia, Ulmus pumila, Salix babylonica, and Melia azedarach, collected from 1963 to 2009 at 47 Chinese Phenological Observation Network (CPON) stations spread across China (from 21° to 50° N). The results of this study show that changes in temperatures in the range of 39-43 days preceding the date of FLD of these plants affected annual variations in FLD, while annual variations in temperature in the range of 71-85 days preceding LCD of these plants affected the date of LCD. Average temperature sensitivity of FLD and LCD for these plants was -3.93 to 3.30 days °C(-1) and 2.11 to 4.43 days °C6301, respectively. Temperature sensitivity of FLD was found to be stronger at lower latitudes or altitude as well as in more continental climates, while the response of LCD showed no consistent pattern. Within the context of significant warming across China during the study period, FLD was found to have advanced by 5.44 days from 1960 to 2009; over the same period, LCD was found to have been delayed by 4.56 days. These findings indicate that the length of the growing season of the four plant species studied was extended by a total of 10.00 days from 1960 to 2009. They also indicate that phenological response to climate is highly heterogeneous spatially.

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[22]
Wang H J, Ge Q S, Dai J H, et al.Geographical pattern in first bloom variability and its relation to temperature sensitivity in the USA and China. International Journal of Biometeorology, 2015, 59(8): 961-969.Advance in spring plant phenology over the last several decades has been found in all continents of the Northern Hemisphere. Compared to the studies detecting phenological trends, the studies investigating the geographical pattern of phenological variability (including mean date and magnitude of variability) are rather limited. In this study, we analyzed spatial pattern of mean date and standard deviation (SD) of first bloom date (FBD) time series (>= 15 years) for black locust (Robinia pseudoacacia) at 22 stations in China, common lilac (Syringa vulgaris) at 79 stations in the Western US and Chinese lilac (Syringa chinensis) at 45 stations in the Eastern US. Subsequently, the impact of geographical factors (latitude, longitude, and altitude) on the mean date and SD was quantified by using the multiple regression analysis method. Meanwhile, the relationship between FBD variability and temperature sensitivity of FBD was examined. Results showed that the mean FBD highly depended on geographical factors for all the three species. Compared to the mean date, the dependence of SD of FBD time series on geographical factors was weaker. The geographical factors could only explain 13 to 31 % of spatial variance in SD of FBD. The negative regression coefficients of latitude (P<0.05 except black locust) indicated that FBD is more variable at lower latitude. At most of stations, significant and negative correlations between FBD and preseason temperature on interannual scale were found, but the temperature sensitivity varied among different stations. The magnitude of temperature sensitivity decreased with increasing latitude. In general, the locations at lower latitude had earlier and more variable spring phenophase and showed stronger phenological response to climate change than the locations at higher latitude.

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[23]
徐韵佳, 戴君虎, 王焕炯, 等. 1985-2012年哈尔滨自然历主要物候期变动特征及对气温变化的响应. 地理研究, 2015, 34(9): 1662-1674.lt;p>利用中国物候观测网观测数据,新编制了哈尔滨地区1985-2012年的自然历。通过与原自然历（1963-1984年）比较,揭示了近30年以来哈尔滨地区21个植物99个物候期的变化特征,并通过物候期与气温的相关分析探讨了物候变化原因。结果表明：自1985年以来,哈尔滨的春季、夏季、秋季的物候期开始日期提前,冬季开始日期推迟。其中春季（以白榆叶芽膨大期为代表）、夏季（以暴马丁香开花始期为代表）、秋季（以金银忍冬果实成熟期为代表）分别提前了7天、6天和19天,冬季（以胡桃楸落叶末期为代表）推迟了2天。各物候期在春季、夏季、秋季的平均日期相较于原自然历提前了3~11天,在冬季推迟了3天。四季各物候期最早日期均以提前为主,夏冬季物候期最晚日期有所推迟。另外,各季节内部分物候期出现的先后次序发生了变化。近30年该地区气温的升高是物候季节开始日期提前的首要原因。且不同植物和物候期对气温变化的响应敏感性不同可解释物候季节内物候期先后次序的变化。</p>

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[Xu Yunjia, Dai Junhu, Wang Huanjiong, et al.Variations of main phenophases of natural calendar and analysis of responses to climate change in Harbin in 1985-2012. Geographical Research, 2015, 34(9): 1662-1674.]

[24]
Zhang H, Yuan W, Liu S, et al.Sensitivity of flowering phenology to changing temperature in China. Journal of Geophysical Research-Biogeosciences, 2015, 120(8): 1658-1665.Plant phenology is one of the preferred indicators of climate change, and its variation potentially impacts community dynamics and ecosystem functions. To better understand the responses of plants' flowering phenology to rising temperatures, we investigated the temperature sensitivity (expressed as the date of changes in phenology per change in temperature in degree Celsius, d C) of flowering phenology for more than 220 plant species at 59 sites in China during the period 1963-1988. Our results indicated that most flowerings in China were significantly sensitive to the temperature in the 2 months (60 days) prior to the flowering dates. Plants in warmer regions showed larger sensitivities to increased temperatures. Species flowering in the late spring and early summer were generally less sensitive to changing temperature than species flowering at other times of the year. For plants flowering in the spring, species that flower earlier showed higher temperature sensitivity; however, for plants flowering in the summer and autumn, species that flower earlier showed lower temperature sensitivity. The responses of the first and last flowering times to changing temperature were mostly consistent, so flowering durations were rarely (6.1%) sensitive to changing temperature. We hypothesize that plants in cold regions may have adapted to the more variable temperatures and thus showed lower temperature sensitivities than plants in warm regions. Overall, the responses of flowering phenology to temperature varied significantly among temperature zones and plant species, so it should be considered carefully when estimating the impacts of climate warming on the terrestrial biosphere.

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[25]
Bai J, Ge Q S, Dai J H.The response of first flowering dates to abrupt climate change in Beijing. Advances in Atmospheric Sciences, 2011, 28(3): 564-572.Abstract Phenological data on the First Flowering Date (FFD) of woody plants in Beijing from 1963–2007 are analyzed. The correlation between each species’ yearly FFD and the mean monthly temperatures for every year over a 45-year period is used to identify the month in which temperature has the most effect on FFD. Through further analysis, the FFDs of 48 woody plant species are shown to have advanced an average of 5.4 days from 1990–2007 compared to 1963–1989. The results indicate that 70.8% of species flowered significantly earlier (7 days on average) during the period 1990–2007, while only one species (2.1%) flowered significantly later. Moreover, the responses of FFD to climate change are shown to be different in two climatic stages, defined by an abrupt climate change point. Thirty-three species which first flower in March and April are sensitive to temperature are examined. The correlation coefficients between FFD and temperature for 20 species during the latter period (1990–2007) are shown to be larger than during the former period (1963–1989), with a difference of around 610.87 days per 1°C on average. The paper concludes that with the warming of climate, the linear trend of FFD variation, as well as its responsiveness to temperature, became more prominent during 1990–2007 than 1963–1989. The data analyzed in this study present a strong biological indicator of climate change in Beijing, and provide further confirmation of previous results from regional and local studies across the Northern Hemisphere. Phenophase variations indicate that the climate is changing rapidly. Keywordsphenology–climate change–first flowering date (FFD)–woody plants

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[26]
Ge Q S, Dai J H, Zheng J Y, et al.Advances in first bloom dates and increased occurrences of yearly second blooms in eastern China since the 1960s: Further phenological evidence of climate warming. Ecological Research, 2011, 26(4): 713-723.Phenology; First bloom; Second bloom; Climate change; China; phenophases; patterns; impacts; plants; onset

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[27]
Zhao J, Zhang Y, Song F, et al.Phenological response of tropical plants to regional climate change in Xishuangbanna, South-western China. Journal of Tropical Ecology, 2013, 29(2): 161-172.ABSTRACT The phenology of temperate plants is vulnerable to climate change. Yet, the phenological responses of tropical plants to climate change are still unclear. In this study, temporal trends (1973–1999) of four phenological events (budburst, growing season, flowering and flowering duration) were studied among 21 plant species in Xishuangbanna Tropical Botanical Garden (south-western China). Fourteen species (67%) showed significant phenological trends during the study period. Seven species (33%) presented delaying trends in budburst (average 1.4 d y611) and such trend was more likely to be presented in those that started budburst earlier in the dry season. Four species (19%) showed trends of extension in growing season (average of 3.5 d y611). These vegetative events appeared to be mainly influenced by increasing temperature. Rainfall showed little effects directly, however, the effects of temperature seemed to largely depend on the moisture condition. Flowering duration of five species (24%) was shortened by average 2.1 d y611 which was most likely to be the result of the decline in sunshine duration during the rainy season. Our results suggest that the phenology of tropical plants has changed significantly in response to the regional climate change but these reactions are somewhat different from those of temperate plants.

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[28]
Bawa K S, Kang H S, Grayum M H.Relationships among time, frequency, and duration of flowering in tropical rain forest trees. American Journal of Botany, 2003, 90(6): 877-887.

[29]
李军, 杨秋珍, 杨康民. 银桂初花物候期的气候条件. 植物生态学报, 2006, 30(3): 421-425.根据江苏省苏州市东山县1956~1984年和1999~2003年银桂(Osmanthusfragrans)物候资料和同期气象资料,应用数理统计方法分析了银桂初花的气候指标,发现银桂初花期与初花前最低气温和相对湿度关系密切。早银桂初花的气候指标为:日最低气温10d滑动平均稳定≤23.0℃,同时日最低气温22.0℃,满足这指标之后3~12d早银桂初花;或当日最低气温10d滑动平均稳定在23.0~25.5℃,同时日平均相对湿度≥88%连续在3d(包括3d)以上,在这日期之后的12d内早银桂初花。晚银桂初花的气候指标为:当日最低气温9d滑动平均稳定≤20℃,同时日最低气温19.5℃,满足这指标后4~12d晚银桂初花;或当日最低气温9d滑动平均稳定在20.0~21.5℃,同时日平均相对湿度≥88%连续在3d(包括3d)以上,在这日期之后的6d内晚银桂初花。日最低气温10d滑动平均稳定25.5℃时,早银桂不开花;日最低气温9d滑动平均稳定21.5℃时,晚银桂不开花。晚银桂和早银桂初花期存在显著的相关,相关系数为0.4438(n=34),平均初花期晚银桂比早银桂迟15d。由于受全球气候变暖的影响,1999~2003年苏州东山地区银桂类初花期比1956~1984年推迟了7~8d。

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[Li Jun, Yang Qiuzhen, Yang Kangmin.Climatic indices for initial flowering in Osmanthus Fragrans. Journal of Plant Ecology, 2006, 30(3): 421-425.]

[30]
Pau S, Wolkovich E M, Cook B I, et al.Predicting phenology by integrating ecology, evolution and climate science. Global Change Biology, 2011, 17(12): 3633-3643.Forecasting how species and ecosystems will respond to climate change has been a major aim of ecology in recent years. Much of this research has focused on phenology – the timing of life‐history events. Phenology has well‐demonstrated links to climate, from genetic to landscape scales; yet our ability to explain and predict variation in phenology across species, habitats and time remains poor. Here, we outline how merging approaches from ecology, climate science and evolutionary biology can advance research on phenological responses to climate variability. Using insight into seasonal and interannual climate variability combined with niche theory and community phylogenetics, we develop a predictive approach for species’ reponses to changing climate. Our approach predicts that species occupying higher latitudes or the early growing season should be most sensitive to climate and have the most phylogenetically conserved phenologies. We further predict that temperate species will respond to climate change by shifting in time, while tropical species will respond by shifting space, or by evolving. Although we focus here on plant phenology, our approach is broadly applicable to ecological research of plant responses to climate variability.

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