植物生态学报 ›› 2013, Vol. 37 ›› Issue (9): 820-829.DOI: 10.3724/SP.J.1258.2013.00086
李立1,2,*(), 杨佳妮3, 崔凯1, 李正红1, 李根前2, 廖声熙1
收稿日期:
2012-12-26
接受日期:
2013-05-22
出版日期:
2013-12-26
发布日期:
2013-09-02
通讯作者:
李立
作者简介:
*E-mail: lili19661118@126.com基金资助:
LI Li1,2,*(), YANG Jia-Ni3, CUI Kai1, R. Talbot TROTTER4,5, LI Zheng-Hong1, LI Gen-Qian2, LIAO Sheng-Xi1
Received:
2012-12-26
Accepted:
2013-05-22
Online:
2013-12-26
Published:
2013-09-02
Contact:
LI Li
摘要:
以进入生殖阶段的8个天然云南铁杉(Tsuga dumosa)种群为对象, 对开花、结实植株(24个样株)的生长量进行测定, 并采用标准枝法对不同树冠层次与方位的开花及结实数量进行调查, 研究了开花结实规律和对其有主要影响的环境因子。结果表明: 云南铁杉雌、雄球花比为1:2, 雌球花转化为果实的数量仅占28%, 胸径为7 cm (树龄约25年)的植株最早进入开花结实阶段, 胸径为25-30 cm的植株开花结实量最高, 胸径为89 cm (树龄约200年)的植株开花结实最晚; 树冠不同层次与方位上开花结实数量有着明显的差异; 相关性分析得出植株胸径与开花结实量极显著相关(p < 0.01)、与冠幅显著相关, 生境中林分郁闭度与开花结实量、株高、胸径之间均呈负相关关系, 其中与结实量的相关性达显著水平; 主成分分析得出植株开花及结实主要受到温度因子、水分因子、光照因子协同作用的影响。云南铁杉雌雄球花的比例偏雄性、结实周期长、结实方式不经济、灾害性天气的影响是造成该种群逐渐走向濒危的主要原因。因此, 采用人工促进天然更新、建立母树林和种子园以提高种子的质量, 对云南铁杉种群的恢复和保护具有重要意义。
李立, 杨佳妮, 崔凯, 李正红, 李根前, 廖声熙. 云南铁杉开花结实特性及其与环境因子的关系. 植物生态学报, 2013, 37(9): 820-829. DOI: 10.3724/SP.J.1258.2013.00086
LI Li, YANG Jia-Ni, CUI Kai, R. Talbot TROTTER, LI Zheng-Hong, LI Gen-Qian, LIAO Sheng-Xi. Strobili and seed characteristics of Tsuga dumosa and its relationship with environmental factors. Chinese Journal of Plant Ecology, 2013, 37(9): 820-829. DOI: 10.3724/SP.J.1258.2013.00086
样地 SP | 经度 Longitude | 纬度 Latitude | 海拔 H (m) | 郁闭度 CD | 年平均 气温 MAT (℃) | 最冷月平 均气温 MTC (℃) | 最暖月平 均气温 MTW (℃) | 年降水量 AP (mm) | ≥10 ℃年 积温 AT≥10℃(℃) | 年日照 时数 ASH (h) | 相对 湿度 RH (%) |
---|---|---|---|---|---|---|---|---|---|---|---|
1 | 99°48′ E | 26°39′ N | 3 027 | 0.48 | 10.8 | 3.1 | 15.5 | 1 351.9 | 3 046.7 | 2 003.6 | 78 |
2 | 99°25′ E | 26°83′ N | 2 840 | 0.62 | 14.7 | 3.8 | 18.2 | 1 066.5 | 3 371.6 | 2 196.8 | 74 |
3 | 99°18′ E | 26°27′ N | 2 956 | 0.55 | 12.1 | 3.6 | 17.2 | 1 022.3 | 3 211.0 | 2 176.4 | 73 |
4 | 100°42′ E | 24°29′ N | 2 704 | 0.72 | 17.6 | 10.9 | 22.8 | 1 103.5 | 6 398.1 | 2 100.9 | 76 |
5 | 101°37′ E | 24°23′ N | 2 890 | 0.61 | 16.8 | 10.3 | 20.9 | 961.3 | 5 711.4 | 2 133.2 | 71 |
6 | 101°22′ E | 24°11′ N | 2 700 | 0.70 | 17.4 | 10.5 | 21.6 | 952.7 | 5 722.8 | 2 252.4 | 69 |
7 | 100°16′ E | 27°09′ N | 2 925 | 0.59 | 16.3 | 9.7 | 20.1 | 1 121.7 | 4 832.5 | 2 092.6 | 77 |
8 | 98°50′ E | 25°21′ N | 2 810 | 0.64 | 15.6 | 8.1 | 20.9 | 974.2 | 4 929.4 | 2 379.6 | 72 |
表1 各种群的地理位置、林分郁闭度及气候状况
Table 1 Geographical position, forest canopy density and climatic conditions of all populations
样地 SP | 经度 Longitude | 纬度 Latitude | 海拔 H (m) | 郁闭度 CD | 年平均 气温 MAT (℃) | 最冷月平 均气温 MTC (℃) | 最暖月平 均气温 MTW (℃) | 年降水量 AP (mm) | ≥10 ℃年 积温 AT≥10℃(℃) | 年日照 时数 ASH (h) | 相对 湿度 RH (%) |
---|---|---|---|---|---|---|---|---|---|---|---|
1 | 99°48′ E | 26°39′ N | 3 027 | 0.48 | 10.8 | 3.1 | 15.5 | 1 351.9 | 3 046.7 | 2 003.6 | 78 |
2 | 99°25′ E | 26°83′ N | 2 840 | 0.62 | 14.7 | 3.8 | 18.2 | 1 066.5 | 3 371.6 | 2 196.8 | 74 |
3 | 99°18′ E | 26°27′ N | 2 956 | 0.55 | 12.1 | 3.6 | 17.2 | 1 022.3 | 3 211.0 | 2 176.4 | 73 |
4 | 100°42′ E | 24°29′ N | 2 704 | 0.72 | 17.6 | 10.9 | 22.8 | 1 103.5 | 6 398.1 | 2 100.9 | 76 |
5 | 101°37′ E | 24°23′ N | 2 890 | 0.61 | 16.8 | 10.3 | 20.9 | 961.3 | 5 711.4 | 2 133.2 | 71 |
6 | 101°22′ E | 24°11′ N | 2 700 | 0.70 | 17.4 | 10.5 | 21.6 | 952.7 | 5 722.8 | 2 252.4 | 69 |
7 | 100°16′ E | 27°09′ N | 2 925 | 0.59 | 16.3 | 9.7 | 20.1 | 1 121.7 | 4 832.5 | 2 092.6 | 77 |
8 | 98°50′ E | 25°21′ N | 2 810 | 0.64 | 15.6 | 8.1 | 20.9 | 974.2 | 4 929.4 | 2 379.6 | 72 |
样株 ST | 胸径 DBH (cm) | 株高 TH (m) | 冠幅 CB (m2) |
---|---|---|---|
1 | 20.2 | 8.3 | 2.4 |
2 | 30.3 | 10.4 | 2.9 |
3 | 25.0 | 9.5 | 2.6 |
4 | 30.5 | 11.8 | 3.2 |
5 | 20.4 | 8.6 | 2.5 |
6 | 26.1 | 11.1 | 2.8 |
7 | 21.4 | 9.4 | 2.5 |
8 | 31.2 | 11.9 | 3.6 |
9 | 26.7 | 10.6 | 3.0 |
10 | 34.6 | 14.2 | 5.1 |
11 | 29.4 | 12.5 | 4.6 |
12 | 25.3 | 10.3 | 3.9 |
13 | 27.6 | 11.6 | 3.5 |
14 | 32.8 | 13.3 | 4.8 |
15 | 24.3 | 9.8 | 3.4 |
16 | 28.6 | 12.2 | 3.7 |
17 | 33.9 | 13.7 | 5.2 |
18 | 24.8 | 10.2 | 3.3 |
19 | 22.6 | 9.3 | 2.6 |
20 | 31.7 | 12.9 | 4.2 |
21 | 26.9 | 11.4 | 3.1 |
22 | 32.2 | 12.7 | 4.4 |
23 | 23.5 | 9.5 | 2.7 |
24 | 27.3 | 11.2 | 3.8 |
表2 云南铁杉开花、结实植株的生长特征
Table 2 Growth characteristics of strobili and seed of Tsuga dumosa
样株 ST | 胸径 DBH (cm) | 株高 TH (m) | 冠幅 CB (m2) |
---|---|---|---|
1 | 20.2 | 8.3 | 2.4 |
2 | 30.3 | 10.4 | 2.9 |
3 | 25.0 | 9.5 | 2.6 |
4 | 30.5 | 11.8 | 3.2 |
5 | 20.4 | 8.6 | 2.5 |
6 | 26.1 | 11.1 | 2.8 |
7 | 21.4 | 9.4 | 2.5 |
8 | 31.2 | 11.9 | 3.6 |
9 | 26.7 | 10.6 | 3.0 |
10 | 34.6 | 14.2 | 5.1 |
11 | 29.4 | 12.5 | 4.6 |
12 | 25.3 | 10.3 | 3.9 |
13 | 27.6 | 11.6 | 3.5 |
14 | 32.8 | 13.3 | 4.8 |
15 | 24.3 | 9.8 | 3.4 |
16 | 28.6 | 12.2 | 3.7 |
17 | 33.9 | 13.7 | 5.2 |
18 | 24.8 | 10.2 | 3.3 |
19 | 22.6 | 9.3 | 2.6 |
20 | 31.7 | 12.9 | 4.2 |
21 | 26.9 | 11.4 | 3.1 |
22 | 32.2 | 12.7 | 4.4 |
23 | 23.5 | 9.5 | 2.7 |
24 | 27.3 | 11.2 | 3.8 |
图1 云南铁杉花、球果、种子形态及生境。A, 开花及结实植株。B, 雌雄球花(顶生为雌球花, 叶腋生为雄球花)。C, 授粉的雌球花。D, 球果。E, 种子(去掉种翅)。F, 云南铁杉生境。
Fig. 1 Shape of strobili, cone and seed of Tsuga dumosa and its habitat. A, Strobili and seed plant. B, Male and female strobilus (female strobilus in tip-growth; male strobilus with axillary). C, Female strobilus of pollination. D, Cone. E, Seed (clear away seed wing). F, Habitat of Tsuga dumosa.
图2 云南铁杉雌、雄球花及球果数量的径级分布(平均值±标准误差)。
Fig. 2 Classes distribution of number of male and female strobilus, cone of Tsuga dumosa in diameter at breast height (DBH) (mean ± SE).
图3 云南铁杉雌、雄球花和球果数量在树冠不同层次及方位上的空间分布。
Fig. 3 Spatial distribution of number of male and female strobilus, cone of Tsuga dumosa in different canopy layers and orientations.
差异源 Difference source | df | SS | MS | F | F0.05 | F0.01 |
---|---|---|---|---|---|---|
上层 Upside layer | 5 | 5 716.335 | 1 143.267 | 127.353** | 8.754 | 9.932 |
中层 Middle layer | 5 | 2 654.246 | 530.849 | 61.267* | 8.754 | 9.932 |
下层 Under layer | 5 | 957.432 | 191.486 | 20.913 | 8.754 | 9.932 |
东向 East | 5 | 1 431.653 | 286.331 | 32.658 | 8.754 | 9.932 |
南向 South | 5 | 1 214.387 | 242.877 | 27.465 | 8.754 | 9.932 |
西向 West | 5 | 2 485.761 | 497.152 | 58.739* | 8.754 | 9.932 |
北向 North | 5 | 2 159.635 | 431.927 | 54.571* | 8.754 | 9.932 |
表3 云南铁杉树冠不同层次及方位上开花数量的方差分析
Table 3 Analysis of variance of strobili quantity of Tsuga dumosa in different canopy layers and orientations
差异源 Difference source | df | SS | MS | F | F0.05 | F0.01 |
---|---|---|---|---|---|---|
上层 Upside layer | 5 | 5 716.335 | 1 143.267 | 127.353** | 8.754 | 9.932 |
中层 Middle layer | 5 | 2 654.246 | 530.849 | 61.267* | 8.754 | 9.932 |
下层 Under layer | 5 | 957.432 | 191.486 | 20.913 | 8.754 | 9.932 |
东向 East | 5 | 1 431.653 | 286.331 | 32.658 | 8.754 | 9.932 |
南向 South | 5 | 1 214.387 | 242.877 | 27.465 | 8.754 | 9.932 |
西向 West | 5 | 2 485.761 | 497.152 | 58.739* | 8.754 | 9.932 |
北向 North | 5 | 2 159.635 | 431.927 | 54.571* | 8.754 | 9.932 |
层次与方位 Layer and orientation | 平均值 Average ($\overline{X}$) | $\overline{X}$–14 876 | $\overline{X}$–18 651 | $\overline{X}$–19 206 | $\overline{X}$–20 309 | $\overline{X}$–21 098 | $\overline{X}$–22 796 |
---|---|---|---|---|---|---|---|
上层 Upside layer | 41 592 | 26 716** | 22 941** | 22 386** | 21 283** | 20 494** | 18 796** |
中层 Middle layer | 22 796 | 7 920* | 4 145* | 3 590 | 2 487 | 1 698 | |
西向 West | 21 098 | 6 222* | 2 447 | 1 892 | 789 | ||
北向 North | 20 309 | 5 433* | 1 658 | 1 103 | |||
东向 East | 19 206 | 4 330 | 555 | ||||
南向 South | 18 651 | 3 775 | |||||
下层 Under layer | 14 876 |
表4 云南铁杉树冠不同层次及方位上开花数量的多重比较
Table 4 Multiple comparison of strobili quantity of Tsuga dumosa in different canopy layers and orientations
层次与方位 Layer and orientation | 平均值 Average ($\overline{X}$) | $\overline{X}$–14 876 | $\overline{X}$–18 651 | $\overline{X}$–19 206 | $\overline{X}$–20 309 | $\overline{X}$–21 098 | $\overline{X}$–22 796 |
---|---|---|---|---|---|---|---|
上层 Upside layer | 41 592 | 26 716** | 22 941** | 22 386** | 21 283** | 20 494** | 18 796** |
中层 Middle layer | 22 796 | 7 920* | 4 145* | 3 590 | 2 487 | 1 698 | |
西向 West | 21 098 | 6 222* | 2 447 | 1 892 | 789 | ||
北向 North | 20 309 | 5 433* | 1 658 | 1 103 | |||
东向 East | 19 206 | 4 330 | 555 | ||||
南向 South | 18 651 | 3 775 | |||||
下层 Under layer | 14 876 |
差异源 Difference source | df | SS | MS | F | F0.05 | F0.01 |
---|---|---|---|---|---|---|
上层 Upside layer | 5 | 653.426 | 130.685 | 55.418* | 2.749 | 3.861 |
中层 Middle layer | 5 | 1 175.248 | 235.050 | 104.136** | 2.749 | 3.861 |
下层 Under layer | 5 | 296.374 | 59.275 | 12.781 | 2.749 | 3.861 |
东向 East | 5 | 557.642 | 111.528 | 46.394* | 2.749 | 3.861 |
南向 South | 5 | 593.286 | 118.657 | 48.563* | 2.749 | 3.861 |
西向 West | 5 | 325.741 | 65.148 | 17.932 | 2.749 | 3.861 |
北向 North | 5 | 381.579 | 76.316 | 22.647 | 2.749 | 3.861 |
表5 云南铁杉树冠不同层次及方位上结实数量的方差分析
Table 5 Analysis of variance of seed quantity of Tsuga dumosa in different canopy layers and orientations
差异源 Difference source | df | SS | MS | F | F0.05 | F0.01 |
---|---|---|---|---|---|---|
上层 Upside layer | 5 | 653.426 | 130.685 | 55.418* | 2.749 | 3.861 |
中层 Middle layer | 5 | 1 175.248 | 235.050 | 104.136** | 2.749 | 3.861 |
下层 Under layer | 5 | 296.374 | 59.275 | 12.781 | 2.749 | 3.861 |
东向 East | 5 | 557.642 | 111.528 | 46.394* | 2.749 | 3.861 |
南向 South | 5 | 593.286 | 118.657 | 48.563* | 2.749 | 3.861 |
西向 West | 5 | 325.741 | 65.148 | 17.932 | 2.749 | 3.861 |
北向 North | 5 | 381.579 | 76.316 | 22.647 | 2.749 | 3.861 |
层次与方位 Layer and orientation | 平均值 Average ($\overline{X}$) | $\overline{X}$–1 006 | $\overline{X}$–1 100 | $\overline{X}$–1 234 | $\overline{X}$–2 577 | $\overline{X}$–2 691 | $\overline{X}$–2 795 |
---|---|---|---|---|---|---|---|
中层 Middle layer | 3 801 | 2 795** | 2 701** | 2 567** | 1 224 | 1 110 | 1 006 |
上层 Upside layer | 2 795 | 1 789* | 1 695* | 1 561* | 218 | 104 | |
南向 South | 2 691 | 1 685* | 1 591* | 1 457* | 114 | ||
东向 East | 2 577 | 1 571* | 1 477* | 1 343 | |||
北向 North | 1 234 | 228 | 134 | ||||
西向 West | 1 100 | 94 | |||||
下层 Under layer | 1 006 |
表6 云南铁杉树冠不同层次及方位上结实数量的多重比较
Table 6 Multiple comparison of seed quantity of Tsuga dumosa in different canopy layers and orientations
层次与方位 Layer and orientation | 平均值 Average ($\overline{X}$) | $\overline{X}$–1 006 | $\overline{X}$–1 100 | $\overline{X}$–1 234 | $\overline{X}$–2 577 | $\overline{X}$–2 691 | $\overline{X}$–2 795 |
---|---|---|---|---|---|---|---|
中层 Middle layer | 3 801 | 2 795** | 2 701** | 2 567** | 1 224 | 1 110 | 1 006 |
上层 Upside layer | 2 795 | 1 789* | 1 695* | 1 561* | 218 | 104 | |
南向 South | 2 691 | 1 685* | 1 591* | 1 457* | 114 | ||
东向 East | 2 577 | 1 571* | 1 477* | 1 343 | |||
北向 North | 1 234 | 228 | 134 | ||||
西向 West | 1 100 | 94 | |||||
下层 Under layer | 1 006 |
NFS | NMS | NCS | TH | DBH | CB | E | N | H | ||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
NFS | 1.000 | 0.516 | 0.951** | 0.262 | 0.936** | 0.197 | 0.324 | -0.602 | -0.569 | |||||||||
NMS | 1.000 | 0.768 | 0.374 | 0.943** | 0.524 | 0.635 | 0.278 | 0.433 | ||||||||||
NCS | 1.000 | 0.067 | 0.925** | 0.873* | 0.147 | -0.094 | -0.902* | |||||||||||
TH | 1.000 | 0.037 | 0.049 | -0.053 | 0.031 | -0.126 | ||||||||||||
DBH | 1.000 | 0.856* | -0.089 | -0.046 | -0.154 | |||||||||||||
CB | 1.000 | 0.072 | -0.025 | -0.098 | ||||||||||||||
MTC | MTW | MAT | AT≥10℃ | AP | RH | ASH | CD | |||||||||||
NFS | 0.513 | 0.632 | 0.458 | 0.715 | -0.746 | -0.619 | 0.885* | -0.731 | ||||||||||
NMS | 0.648 | 0.724 | 0.605 | 0.793 | -0.681 | -0. 573 | 0.891* | -0.627 | ||||||||||
NCS | 0.846* | 0.851* | 0.863* | 0.957** | 0.948** | 0.837* | 0.674 | -0.829* | ||||||||||
TH | -0.710 | 0.203 | 0.114 | 0.074 | 0.054 | 0.092 | 0.380 | -0.016 | ||||||||||
DBH | -0.345 | 0.428 | 0.309 | 0.831* | 0.876* | 0.704 | 0.538 | -0.064 | ||||||||||
CB | -0.184 | 0.317 | 0.256 | 0.312 | 0.167 | 0.068 | 0.457 | 0.273 |
表7 云南铁杉开花结实量和植株生长量间及其与环境因子间的相关系数
Table 7 Correlation coefficient between strobili and seed quantity, growth increment and environmental factors of Tsuga dumosa
NFS | NMS | NCS | TH | DBH | CB | E | N | H | ||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
NFS | 1.000 | 0.516 | 0.951** | 0.262 | 0.936** | 0.197 | 0.324 | -0.602 | -0.569 | |||||||||
NMS | 1.000 | 0.768 | 0.374 | 0.943** | 0.524 | 0.635 | 0.278 | 0.433 | ||||||||||
NCS | 1.000 | 0.067 | 0.925** | 0.873* | 0.147 | -0.094 | -0.902* | |||||||||||
TH | 1.000 | 0.037 | 0.049 | -0.053 | 0.031 | -0.126 | ||||||||||||
DBH | 1.000 | 0.856* | -0.089 | -0.046 | -0.154 | |||||||||||||
CB | 1.000 | 0.072 | -0.025 | -0.098 | ||||||||||||||
MTC | MTW | MAT | AT≥10℃ | AP | RH | ASH | CD | |||||||||||
NFS | 0.513 | 0.632 | 0.458 | 0.715 | -0.746 | -0.619 | 0.885* | -0.731 | ||||||||||
NMS | 0.648 | 0.724 | 0.605 | 0.793 | -0.681 | -0. 573 | 0.891* | -0.627 | ||||||||||
NCS | 0.846* | 0.851* | 0.863* | 0.957** | 0.948** | 0.837* | 0.674 | -0.829* | ||||||||||
TH | -0.710 | 0.203 | 0.114 | 0.074 | 0.054 | 0.092 | 0.380 | -0.016 | ||||||||||
DBH | -0.345 | 0.428 | 0.309 | 0.831* | 0.876* | 0.704 | 0.538 | -0.064 | ||||||||||
CB | -0.184 | 0.317 | 0.256 | 0.312 | 0.167 | 0.068 | 0.457 | 0.273 |
变量 Variable | 主成分 Principal component | ||
---|---|---|---|
1 | 2 | 3 | |
E | 0.293 | 0.628 | 0.532 |
N | 0.267 | 0.451 | 0.674 |
H | -0.018 | 0.236 | 0.095 |
MTC | 0.931 | -0.160 | 0.013 |
MTW | 0.956 | -0.122 | 0.085 |
MAT | 0.942 | -0.143 | 0.056 |
AT≥10℃ | 0.984 | -0.097 | 0.132 |
AP | -0.165 | 0.928 | -0.073 |
RH | -0.240 | 0.745 | -0.327 |
ASH | 0.021 | -0.252 | 0.896 |
特征值 Eigenvalue | 7.423 | 3.054 | 2.254 |
方差贡献率 % of variance | 52.367 | 21.246 | 8.132 |
累积贡献率 % of cumulative | 52.367 | 73.613 | 81.745 |
表8 前3个主成分因子指标负荷量
Table 8 First three principal component loadfactors of index
变量 Variable | 主成分 Principal component | ||
---|---|---|---|
1 | 2 | 3 | |
E | 0.293 | 0.628 | 0.532 |
N | 0.267 | 0.451 | 0.674 |
H | -0.018 | 0.236 | 0.095 |
MTC | 0.931 | -0.160 | 0.013 |
MTW | 0.956 | -0.122 | 0.085 |
MAT | 0.942 | -0.143 | 0.056 |
AT≥10℃ | 0.984 | -0.097 | 0.132 |
AP | -0.165 | 0.928 | -0.073 |
RH | -0.240 | 0.745 | -0.327 |
ASH | 0.021 | -0.252 | 0.896 |
特征值 Eigenvalue | 7.423 | 3.054 | 2.254 |
方差贡献率 % of variance | 52.367 | 21.246 | 8.132 |
累积贡献率 % of cumulative | 52.367 | 73.613 | 81.745 |
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