内蒙古典型草原植物叶片碳氮磷化学计量特征的季节动态

doi:10.17521/cjpe.2020.0105

植物生态学报 ›› 2020, Vol. 44 ›› Issue (11): 1138-1153.DOI: 10.17521/cjpe.2020.0105

所属专题：生态化学计量

内蒙古典型草原植物叶片碳氮磷化学计量特征的季节动态

熊星烁¹^,², 蔡宏宇², 李耀琪², 马文红³, 牛克昌⁴, 陈迪马⁵, 刘娜娜², 苏香燕², 景鹤影², 冯晓娟⁶, 曾辉¹^,²^,^*(), 王志恒²^,^*()

¹北京大学深圳研究生院城市规划与设计学院, 广东深圳 518055
²北京大学生态研究中心, 北京大学城市与环境学院, 北京大学地表过程分析与模拟教育部重点实验室, 北京 100871
³内蒙古大学生态与环境学院, 呼和浩特 010021
⁴南京大学生命科学学院, 南京 210023
⁵三峡大学生物与制药学院, 湖北宜昌 443002
⁶中国科学院植物研究所植被与环境变化国家重点实验室, 北京 100093

收稿日期:2020-04-15 接受日期:2020-07-16 出版日期:2020-11-20 发布日期:2021-01-05
通讯作者: 曾辉,王志恒
作者简介:王志恒: ORCID:0000-0003-0808-7780,Wang ZH: zhiheng.wang@pku.edu.cn
*Zeng H: zengh@pkusz.edu.cn;
基金资助:
国家重点研发计划(2018YFA0606104);国家自然科学基金(31988102)

Seasonal dynamics of leaf C, N and P stoichiometry in plants of typical steppe in Nei Mongol, China

XIONG Xing-Shuo¹^,², CAI Hong-Yu², LI Yao-Qi², MA Wen-Hong³, NIU Ke-Chang⁴, CHEN Di-Ma⁵, LIU Na-Na², SU Xiang-Yan², JING He-Ying², FENG Xiao-Juan⁶, ZENG Hui¹^,²^,^*(), WANG Zhi-Heng²^,^*()

¹School of Urban Planning and Design, Peking University Shenzhen Graduate School, Shenzhen, Guangdong 518055, China
²Institute of Ecology, Key Laboratory for Earth Surface Processes of the Ministry of Education, College of Urban and Environmental Sciences, Peking University, Beijing 100871, China
³School of Ecology and Environment, Inner Mongolia University, Hohhot 010021, China
⁴School of Life Sciences, Nanjing University, Nanjing 210023, China
⁵College of Biology and Pharmacy, China Three Gorges University, Yichang, Hubei 443002, China
⁶State Key Laboratory of Vegetation and Environmental Change, Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China

Received:2020-04-15 Accepted:2020-07-16 Online:2020-11-20 Published:2021-01-05
Contact: ZENG Hui,WANG Zhi-Heng
Supported by:
National Key R&D Program of China(2018YFA0606104);National Natural Science Foundation of China(31988102)

摘要/Abstract

摘要：

分析植物叶片(C)、氮(N)、磷(P)含量及其比值的季节动态, 不仅有助于认识植物生长发育和养分吸收利用等生理生态过程, 也有利于认识植物化学计量的动态平衡关系。该文选择内蒙古典型温带草原18种常见植物, 在生长季的6-9月, 每半月一次进行连续采样, 在此基础上分析了叶片C、N、P含量及其比值在生长季内的变化。主要结果: 1)植物叶片C、N、P含量及其比值的季节性变化在不同功能类群间不同步, 其中叶片N、P含量的季节变化体现了明显的稀释作用。2)叶片C、N、P含量及其比值在不同功能类群间差异显著, 单子叶、多年生禾草类的叶片N、P含量显著低于双子叶和多年生杂类草植物, 而其叶片C:N、C:P则高于双子叶和多年生杂类草植物。3)叶片N、P含量显著正相关, 叶片C:N和C:P分别与N和P含量显著负相关, 可能体现了植物体内营养元素间的内在耦合机制。4)叶片N含量与C:N, 叶片P含量与C:P以及叶片N含量与P含量均呈现等速生长关系, 且等速生长关系在生长季保持稳定。

关键词: 季节动态, 生态化学计量学, 异速生长, 叶片, 典型草原

Abstract:

Aims Exploring the seasonal dynamics in leaf carbon (C), nitrogen (N) and phosphorus (P) concentrations and their ecological stoichiometric characteristics will enhance our understanding about physiological and ecological processes such as plant growth and development and nutrient uptake and utilization as well as dynamic equilibrium relationship among plant stoichiometry.
Methods Here, we collected leaf samples of 18 dominant plant species semimonthly through growing season (i.e. from June 2nd to Sept. 2nd) from a long-term fenced site in a typical steppe in Xilinhot of Nei Mongol, China. Leaf C, N and P concentrations were measured. Seasonal changes in leaf C, N and P concentrations and their ratios were explored and their differences between different species groups were analyzed using one-way ANOVA. The relationships between leaf C, N and P concentrations and their ratios were analyzed using correlation analysis. Lastly, the allometric relationships between the concentrations of different elements were analyzed using Standardized Major Axis.
Important findings Seasonal trends in leaf C, N and P concentrations and their ratios were not consistent with each other and also differed between different functional groups. Specifically, the variation of leaf N and P concentrations for all functional groups showed obvious dilution effect. Monocotyledons and perennial grasses had lower leaf N and P concentrations but much higher leaf C:N and C:P mass ratio than dicotyledons and perennial forbs, respectively. Leaf N concentration was positively correlated with leaf P concentration while leaf C:N and C:P mass ratios were negatively correlated with leaf N and P concentrations respectively, indicating the internal coupling mechanism between nutrient elements in plants. Allometric analyses showed that leaf N concentration and C:N mass ratio, leaf P concentration and C:P mass ratio as well as leaf N and P concentrations all maintained the same growth rate respectively among species through most time of growing season.

Key words: seasonal dynamic, stoichiometry, allometry, leaf, typical steppe

熊星烁, 蔡宏宇, 李耀琪, 马文红, 牛克昌, 陈迪马, 刘娜娜, 苏香燕, 景鹤影, 冯晓娟, 曾辉, 王志恒. 内蒙古典型草原植物叶片碳氮磷化学计量特征的季节动态. 植物生态学报, 2020, 44(11): 1138-1153. DOI: 10.17521/cjpe.2020.0105

XIONG Xing-Shuo, CAI Hong-Yu, LI Yao-Qi, MA Wen-Hong, NIU Ke-Chang, CHEN Di-Ma, LIU Na-Na, SU Xiang-Yan, JING He-Ying, FENG Xiao-Juan, ZENG Hui, WANG Zhi-Heng. Seasonal dynamics of leaf C, N and P stoichiometry in plants of typical steppe in Nei Mongol, China. Chinese Journal of Plant Ecology, 2020, 44(11): 1138-1153. DOI: 10.17521/cjpe.2020.0105

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物种 Species	发育类群 Phylogenetic group	生活型 Life form	6月 June	7月 July	8月 Aug.	9月 Sept.
羽茅 Achnatherum sibiricum	单子叶 Monocotyledons	多年生禾草 Perennial grasses	A	A	B, C	D
长柱沙参 Adenophora stenanthina	双子叶 Dicotyledons	多年生杂类草 Perennial forbs	A	A, B	B	B, C
冰草 Agropyron cristatum	单子叶 Monocotyledons	多年生禾草 Perennial grasses	A	B, C	D	D, E
矮韭 Allium anisopodium	单子叶 Monocotyledons	多年生杂类草 Perennial forbs	A, B	C	D	D
黄花葱 Allium condensatum	单子叶 Monocotyledons	多年生杂类草 Perennial forbs	A, B	C	D	D
山韭 Allium senescens	单子叶 Monocotyledons	多年生杂类草 Perennial forbs	A, B	C	D	D
细叶韭 Allium tenuissimum	单子叶 Monocotyledons	多年生杂类草 Perennial forbs	A, B	C	D	D
小叶锦鸡儿 Caragana microphylla	双子叶 Dicotyledons	-	B	C	C	D
糙隐子草 Cleistogenes squarrosa	单子叶 Monocotyledons	多年生禾草 Perennial grasses	A	A	B, C, D	D, E
狗娃花 Aster hispidus	双子叶 Dicotyledons	多年生杂类草 Perennial forbs	B, C	C	C, D	D, E
木地肤 Kochia prostrata	双子叶 Dicotyledons	-	A	B, C	C, D	D, E
[艹/洽]草 Koeleria macrantha	单子叶 Monocotyledons	多年生禾草 Perennial grasses	B, C	D	D	D, E
羊草 Leymus chinensis	单子叶 Monocotyledons	多年生禾草 Perennial grasses	A	B, C	C, D	D, E
裂叶荆芥 Nepeta tenuifolia	双子叶 Dicotyledons	多年生杂类草 Perennial forbs	A	B, C	C	D
二裂委陵菜 Potentilla bifurca	双子叶 Dicotyledons	多年生杂类草 Perennial forbs	A, B	B, C	C, D	D, E
麻花头 Klasea centauroides	双子叶 Dicotyledons	多年生杂类草 Perennial forbs	B, C	C	C, D	D
大针茅 Stipa grandis	单子叶 Monocotyledons	多年生禾草 Perennial grasses	A	A	B, C	C, D
展枝唐松草 Thalictrum squarrosum	双子叶 Dicotyledons	多年生杂类草 Perennial forbs	A	B, C	C	D

物种 Species	发育类群 Phylogenetic group	生活型 Life form	6月 June	7月 July	8月 Aug.	9月 Sept.
羽茅 Achnatherum sibiricum	单子叶 Monocotyledons	多年生禾草 Perennial grasses	A	A	B, C	D
长柱沙参 Adenophora stenanthina	双子叶 Dicotyledons	多年生杂类草 Perennial forbs	A	A, B	B	B, C
冰草 Agropyron cristatum	单子叶 Monocotyledons	多年生禾草 Perennial grasses	A	B, C	D	D, E
矮韭 Allium anisopodium	单子叶 Monocotyledons	多年生杂类草 Perennial forbs	A, B	C	D	D
黄花葱 Allium condensatum	单子叶 Monocotyledons	多年生杂类草 Perennial forbs	A, B	C	D	D
山韭 Allium senescens	单子叶 Monocotyledons	多年生杂类草 Perennial forbs	A, B	C	D	D
细叶韭 Allium tenuissimum	单子叶 Monocotyledons	多年生杂类草 Perennial forbs	A, B	C	D	D
小叶锦鸡儿 Caragana microphylla	双子叶 Dicotyledons	-	B	C	C	D
糙隐子草 Cleistogenes squarrosa	单子叶 Monocotyledons	多年生禾草 Perennial grasses	A	A	B, C, D	D, E
狗娃花 Aster hispidus	双子叶 Dicotyledons	多年生杂类草 Perennial forbs	B, C	C	C, D	D, E
木地肤 Kochia prostrata	双子叶 Dicotyledons	-	A	B, C	C, D	D, E
[艹/洽]草 Koeleria macrantha	单子叶 Monocotyledons	多年生禾草 Perennial grasses	B, C	D	D	D, E
羊草 Leymus chinensis	单子叶 Monocotyledons	多年生禾草 Perennial grasses	A	B, C	C, D	D, E
裂叶荆芥 Nepeta tenuifolia	双子叶 Dicotyledons	多年生杂类草 Perennial forbs	A	B, C	C	D
二裂委陵菜 Potentilla bifurca	双子叶 Dicotyledons	多年生杂类草 Perennial forbs	A, B	B, C	C, D	D, E
麻花头 Klasea centauroides	双子叶 Dicotyledons	多年生杂类草 Perennial forbs	B, C	C	C, D	D
大针茅 Stipa grandis	单子叶 Monocotyledons	多年生禾草 Perennial grasses	A	A	B, C	C, D
展枝唐松草 Thalictrum squarrosum	双子叶 Dicotyledons	多年生杂类草 Perennial forbs	A	B, C	C	D

		功能类群 Functional group	样本数 Sample number	平均值 Mean	最小值 Minimum	最大值 Maximum	标准偏差 Standard deviation	变异系数 Coefficient of variation
碳含量 C concentration (mg·g^-1)	发育类群 Phylogenetic group	单子叶 Monocotyledons	70	431.87^a	391.36	468.83	18.2	0.04
	发育类群 Phylogenetic group	双子叶 Dicotyledons	56	435.43^a	369.88	474.75	21.03	0.05
	生活型 Life form	多年生禾草 Perennial grasses	42	443.66^a	413.25	468.83	11.36	0.03
	生活型 Life form	多年生杂类草 Perennial forbs	70	429.13^b	390.53	474.75	19.89	0.05
	全部 All		126	433.45	369.88	474.75	19.51	0.05
氮含量 N concentration (mg·g^-1)	发育类群 Phylogenetic group	单子叶 Monocotyledons	70	24.69^b	15.30	48.14	6.35	0.26
	发育类群 Phylogenetic group	双子叶 Dicotyledons	56	32.33^a	20.90	53.79	8.29	0.26
	生活型 Life form	多年生禾草 Perennial grasses	42	23.50^b	15.30	35.25	5.13	0.22
	生活型 Life form	多年生杂类草 Perennial forbs	70	29.85^a	17.00	53.79	8.42	0.28
	全部 All		126	28.08	15.30	53.79	8.19	0.29
磷含量 P concentration (mg·g^-1)	发育类群 Phylogenetic group	单子叶 Monocotyledons	70	1.41^b	0.80	3.09	0.44	0.31
	发育类群 Phylogenetic group	双子叶 Dicotyledons	56	1.75^a	1.04	3.60	0.49	0.28
	生活型 Life form	多年生禾草 Perennial grasses	42	1.22^b	0.80	1.99	0.25	0.20
	生活型 Life form	多年生杂类草 Perennial forbs	70	1.75^a	1.08	3.60	0.51	0.29
	全部 All		126	1.56	0.80	3.60	0.49	0.31
C:N	发育类群 Phylogenetic group	单子叶 Monocotyledons	70	18.55^a	8.48	30.23	4.43	0.24
	发育类群 Phylogenetic group	双子叶 Dicotyledons	56	14.32^b	8.33	21.85	3.59	0.25
	生活型 Life form	多年生禾草 Perennial grasses	42	19.76^a	12.36	30.23	4.28	0.22
	生活型 Life form	多年生杂类草 Perennial forbs	70	15.44^b	8.33	23.09	3.99	0.26
	全部 All		126	16.67	8.33	30.23	4.58	0.27
C:P	发育类群 Phylogenetic group	单子叶 Monocotyledons	70	330.93^a	130.66	573.06	91.16	0.28
	发育类群 Phylogenetic group	双子叶 Dicotyledons	56	267.19^b	118.13	423.14	70.72	0.26
	生活型 Life form	多年生禾草 Perennial grasses	42	378.07^a	218.89	573.06	74.78	0.20
	生活型 Life form	多年生杂类草 Perennial forbs	70	262.79^b	118.13	423.14	68.44	0.26
	全部 All		126	302.60	118.13	573.06	88.31	0.29
N:P	发育类群 Phylogenetic group	单子叶 Monocotyledons	70	17.95^a	12.81	29.33	3.52	0.20
	发育类群 Phylogenetic group	双子叶 Dicotyledons	56	18.78^a	13.76	26.99	2.80	0.15
	生活型 Life form	多年生禾草 Perennial grasses	42	19.50^a	14.26	29.33	3.59	0.18
	生活型 Life form	多年生杂类草 Perennial forbs	70	17.16^b	12.81	23.62	2.34	0.14
	全部 All		126	18.32	12.81	29.33	3.24	0.18

		功能类群 Functional group	样本数 Sample number	平均值 Mean	最小值 Minimum	最大值 Maximum	标准偏差 Standard deviation	变异系数 Coefficient of variation
碳含量 C concentration (mg·g^-1)	发育类群 Phylogenetic group	单子叶 Monocotyledons	70	431.87^a	391.36	468.83	18.2	0.04
	发育类群 Phylogenetic group	双子叶 Dicotyledons	56	435.43^a	369.88	474.75	21.03	0.05
	生活型 Life form	多年生禾草 Perennial grasses	42	443.66^a	413.25	468.83	11.36	0.03
	生活型 Life form	多年生杂类草 Perennial forbs	70	429.13^b	390.53	474.75	19.89	0.05
	全部 All		126	433.45	369.88	474.75	19.51	0.05
氮含量 N concentration (mg·g^-1)	发育类群 Phylogenetic group	单子叶 Monocotyledons	70	24.69^b	15.30	48.14	6.35	0.26
	发育类群 Phylogenetic group	双子叶 Dicotyledons	56	32.33^a	20.90	53.79	8.29	0.26
	生活型 Life form	多年生禾草 Perennial grasses	42	23.50^b	15.30	35.25	5.13	0.22
	生活型 Life form	多年生杂类草 Perennial forbs	70	29.85^a	17.00	53.79	8.42	0.28
	全部 All		126	28.08	15.30	53.79	8.19	0.29
磷含量 P concentration (mg·g^-1)	发育类群 Phylogenetic group	单子叶 Monocotyledons	70	1.41^b	0.80	3.09	0.44	0.31
	发育类群 Phylogenetic group	双子叶 Dicotyledons	56	1.75^a	1.04	3.60	0.49	0.28
	生活型 Life form	多年生禾草 Perennial grasses	42	1.22^b	0.80	1.99	0.25	0.20
	生活型 Life form	多年生杂类草 Perennial forbs	70	1.75^a	1.08	3.60	0.51	0.29
	全部 All		126	1.56	0.80	3.60	0.49	0.31
C:N	发育类群 Phylogenetic group	单子叶 Monocotyledons	70	18.55^a	8.48	30.23	4.43	0.24
	发育类群 Phylogenetic group	双子叶 Dicotyledons	56	14.32^b	8.33	21.85	3.59	0.25
	生活型 Life form	多年生禾草 Perennial grasses	42	19.76^a	12.36	30.23	4.28	0.22
	生活型 Life form	多年生杂类草 Perennial forbs	70	15.44^b	8.33	23.09	3.99	0.26
	全部 All		126	16.67	8.33	30.23	4.58	0.27
C:P	发育类群 Phylogenetic group	单子叶 Monocotyledons	70	330.93^a	130.66	573.06	91.16	0.28
	发育类群 Phylogenetic group	双子叶 Dicotyledons	56	267.19^b	118.13	423.14	70.72	0.26
	生活型 Life form	多年生禾草 Perennial grasses	42	378.07^a	218.89	573.06	74.78	0.20
	生活型 Life form	多年生杂类草 Perennial forbs	70	262.79^b	118.13	423.14	68.44	0.26
	全部 All		126	302.60	118.13	573.06	88.31	0.29
N:P	发育类群 Phylogenetic group	单子叶 Monocotyledons	70	17.95^a	12.81	29.33	3.52	0.20
	发育类群 Phylogenetic group	双子叶 Dicotyledons	56	18.78^a	13.76	26.99	2.80	0.15
	生活型 Life form	多年生禾草 Perennial grasses	42	19.50^a	14.26	29.33	3.59	0.18
	生活型 Life form	多年生杂类草 Perennial forbs	70	17.16^b	12.81	23.62	2.34	0.14
	全部 All		126	18.32	12.81	29.33	3.24	0.18

组别 Group	C:N-N			C:P-P			P-N
组别 Group	斜率 Slope	R²	p	斜率 Slope	R²	p	斜率 Slope	R²	p
06-02	-1.04 (-1.16, -0.93)	0.96	<0.001	-1.04 (-1.11, -0.97)	0.99	<0.001	1.68 (1.26, 2.23)	0.70	<0.001
06-17	-1.00 (-1.10, -0.90)	0.96	<0.001	-1.03 (-1.12, -0.95)	0.97	<0.001	1.16 (0.87, 1.53)	0.71	<0.001
07-02	-0.95 (-1.03, -0.88)	0.98	<0.001	-0.98 (-1.06, -0.90)	0.97	<0.001	0.98 (0.72, 1.32)	0.67	<0.001
07-17	-0.97 (-1.07, -0.87)	0.96	<0.001	-1.05 (-1.15, -0.96)	0.97	<0.001	0.97 (0.68, 1.38)	0.54	<0.001
08-02	-0.97 (-1.05, -0.89)	0.98	<0.001	-1.02 (-1.12, -0.93)	0.97	<0.001	0.90 (0.65, 1.24)	0.62	<0.001
08-17	-1.03 (-1.17, -0.90)	0.94	<0.001	-1.10 (-1.23, -0.98)	0.95	<0.001	1.08 (0.76, 1.54)	0.54	<0.001
09-02	-1.03 (-1.13, -0.93)	0.97	<0.001	-1.08 (-1.16, -1.01)	0.98	<0.001	1.13 (0.80, 1.60)	0.55	<0.001
	共同斜率 Common slope	斜率异质性 Heterogeneity of slopes		共同斜率 Common slope	斜率异质性 Heterogeneity of slopes		共同斜率 Common slope	斜率异质性 Heterogeneity of slopes
全部 All	-0.99 (-1.03, -0.95)	p = 0.77		-1.04 (-1.07, -1.01)	p = 0.58		1.11 (0.97, 1.26)	p = 0.10

内蒙古典型草原植物叶片碳氮磷化学计量特征的季节动态

Seasonal dynamics of leaf C, N and P stoichiometry in plants of typical steppe in Nei Mongol, China

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参数 Parameter	发育类群 Phylogenetic group
参数 Parameter	变异来源 Source of variation	离差平方和 SS	均方 MS	F
碳含量 C concentration (mg·g^-1)	发育类群 Phylogenetic group	396	395.9	1.15
	采样日期² Sampling date²	2 867	2 867.2	8.33^**
	采样日期 Sampling date	1 741	1 740.6	5.06^*
	发育类群×采样日期² Phylogenetic group × Sampling date²	169	168.6	0.49
	发育类群×采样日期 Phylogenetic group × Sampling date	1 088	1 088.4	3.16^?
氮含量 N concentration (mg·g^-1)	发育类群 Phylogenetic group	1 815	1 815.2	61.89^***
	采样日期² Sampling date²	491	490.7	16.73^***
	采样日期 Sampling date	2 280	2 279.7	77.73^***
	发育类群×采样日期² Phylogenetic group × Sampling date²	93	93.1	3.17^?
	发育类群×采样日期 Phylogenetic group × Sampling date	184	183.7	6.26^*
磷含量 P concentration (mg·g^-1)	发育类群 Phylogenetic group	3.48	3.48	22.78^***
	采样日期² Sampling date²	2.18	2.18	14.25^***
	采样日期 Sampling date	4.86	4.86	31.82^***
	发育类群×采样日期² Phylogenetic group × Sampling date²	0.03	0.03	0.20
	发育类群×采样日期 Phylogenetic group × Sampling date	1.40	1.40	9.18^**
C:N	发育类群 Phylogenetic group	554.8	554.8	56.52^***
	采样日期² Sampling date²	135.2	135.2	13.78^***
	采样日期 Sampling date	696.9	696.9	70.80^***
	发育类群×采样日期² Phylogenetic group × Sampling date²	47.8	47.8	4.87^*
	发育类群×采样日期 Phylogenetic group × Sampling date	5.5	5.5	0.56
C:P	发育类群 Phylogenetic group	126 456	126 456	23.29^***
	采样日期²Sampling date²	42 934	42 934	7.91^**
	采样日期 Sampling date	118 984	118 984	21.92^***
	发育类群×采样日期² Phylogenetic group × Sampling date²	5 798	5 798	1.07
	发育类群×采样日期 Phylogenetic group × Sampling date	29 439	29 439	5.42^*
N:P	发育类群 Phylogenetic group	20.9	20.9	2.13
	采样日期² Sampling date²	0.2	0.22	0.02
	采样日期 Sampling date	51.5	51.5	5.26^*
	发育类群×采样日期² Phylogenetic group × Sampling date²	14.7	14.75	1.51
	发育类群×采样日期 Phylogenetic group × Sampling date	46.9	46.9	4.80^*
参数 Parameter	生活型 Life form
参数 Parameter	变异来源 Source of variation	离差平方和 SS	均方 MS	F
碳含量 C concentration (mg·g^-1)	生活型 Life form	5 542	5 542	20.83^***
	采样日期² Sampling date²	2 512	2 512	9.44^**
	采样日期 Sampling date	1 810	1 810	6.80^*
	生活型×采样日期² Life form × Sampling date²	10	10	0.04
	生活型×采样日期 Life form × Sampling date	38	38	0.14
氮含量 N concentration (mg·g^-1)	生活型 Life form	1 060	1 059.6	31.75^***
	采样日期² Sampling date²	526	525.5	15.75^***
	采样日期 Sampling date	1 833	1 832.8	54.92^***
	生活型×采样日期² Life form × Sampling date²	3	3.1	0.09
	生活型×采样日期 Life form × Sampling date	78	78.1	2.34
磷含量 P concentration (mg·g^-1)	生活型 Life form	7.54	7.54	59.98^***
	采样日期² Sampling date²	2.28	2.28	18.17^***
	采样日期 Sampling date	3.53	3.53	28.10^***
	生活型×采样日期² Life form × Sampling date²	0.58	0.58	4.58^*
	生活型×采样日期 Life form × Sampling date	0.76	0.76	6.07^*
C:N	生活型 Life form	490.9	490.9	48.07^***
	采样日期² Sampling date²	149.0	149.0	14.58^***
	采样日期 Sampling date	614.6	614.6	60.17^***
	生活型×采样日期² Life form × Sampling date²	2.1	2.1	0.21
	生活型×采样日期 Life form × Sampling date	3.3	3.3	0.32
C:P	生活型 Life form	348 869	348 869	88.98^***
	采样日期² Sampling date²	49 510	49 510	12.63^***
	采样日期 Sampling date	82 881	82 881	21.14^***
	生活型×采样日期² Life form × Sampling date²	1 157	1 157	0.30
	生活型×采样日期 Life form × Sampling date	3 377	3 377	0.86
N:P	生活型 Life form	143.7	143.74	19.92^***
	采样日期² Sampling date²	0.5	0.46	0.06
	采样日期 Sampling date	73.3	73.3	10.16^**
	生活型×采样日期² Life form × Sampling date²	34.8	34.82	4.83^*
	生活型×采样日期 Life form × Sampling date	32.8	32.80	4.55^*

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