西双版纳热带季节雨林的生物量及其分配特征

doi:10.17521/cjpe.2007.0003

植物生态学报 ›› 2007, Vol. 31 ›› Issue (1): 11-22.DOI: 10.17521/cjpe.2007.0003

西双版纳热带季节雨林的生物量及其分配特征

吕晓涛¹^,²(), 唐建维¹^,^*(), 何有才³, 段文贵³, 宋军平³, 许海龙³, 朱胜忠³

1 中国科学院西双版纳热带植物园,云南勐腊 666303
2 中国科学院研究生院,北京 100049
3 西双版纳国家级自然保护区管理局勐腊管理所,云南勐腊 666300

收稿日期:2005-09-19 接受日期:2006-01-26 出版日期:2007-09-19 发布日期:2007-01-30
通讯作者: 唐建维
作者简介:* tangjw@xtbg.org.cn
基金资助:
中国科学院野外台站基金项目;“西部之光”资助项目和中国科学院知识创新工程重大项目(KZCX1-SW-01)

BIOMASS AND ITS ALLOCATION IN TROPICAL SEASONAL RAIN FOREST IN XISHUANGBANNA, SOUTHWEST CHINA

LÁ Xiao-Tao¹^,²(), TANG Jian-Wei¹^,^*(), HE You-Cai³, DUAN Wen-Gui³, SONG Jun-Ping³, XU Hai-Long³, ZHU Sheng-Zhong³

¹Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Mengla, Yunnan 666303, China,
²Graduate University of Chinese Academy of Sciences, Beijing 100049, China, and
³Mengla Institute of Conservation, Xishuangbanna Bureau of National Nature Reserve, Mengla, Yunnan 666300, China

Received:2005-09-19 Accepted:2006-01-26 Online:2007-09-19 Published:2007-01-30
Contact: TANG Jian-Wei
About author:First author contact:E-mail of the first author: lxtao@xtbg.org.cn.

摘要/Abstract

摘要：

根据3块1 hm² 样地的调查资料,利用123株样木数据建立以胸径(D)为单变量的生物量预测方程。采用样木回归分析法(乔木层、木质藤本)和样方收获法(灌木层、草本层),获取西双版纳热带季节雨林的生物量,并分析了其组成和分配特征。结果表明,西双版纳热带季节雨林的总生物量为(423.908±109.702) Mg·hm^-2(平均值±标准差,n=3),其中活体植物生物量占95.28%,粗死木质残体占4.07%,地上凋落物占 0.64%。在其层次分配方面:乔木层优势明显,占98.09%±0.60%;其次为木质藤本,占0.83%±0.31%;灌木层和草本层生物量均小于木质藤本的生物量;附生植物最低,仅为0.06%±0.03%。总生物量的器官分配以茎所占比例最高,达68.33%;根、枝、叶的比例分别为18.91%、11.07%和1.65%。乔木层生物量的径级分配主要集中于中等径级和最大径级。大树(D>70 cm)具有较高的生物量,占整个乔木层的43.67%±12.67%。树种分配方面,生物量排序前10位的树种占乔木层总生物量的63.43%±4.09%,生物量集中分配于少量优势树种。西双版纳热带季节雨林乔木层叶面积指数为6.39±0.85。西双版纳热带季节雨林乔木层的地上生物量位于世界热带湿润森林的中下范围。

关键词: 生物量, 生物量分配, 回归模型, 热带季节雨林, 西双版纳

Abstract:

Aims Changes in the biomass of tropical forests play an important role in the global carbon cycle, but the biomass of these forests has been poorly quantified. A strategy for regional biomass estimation should supplement previous surveys with new data. Accurate data are necessary for reducing the uncertainty in the carbon budget of tropical regions.

Methods Biomass and its allocation were estimated for the tropical seasonal rain forest in Xishuangbanna, southwest China. Regression models relating tree biomass to DBH (diameter at breast height, 1.3 m) were developed, and a power-law allometric relationship W=aD^b was used to estimate the tree biomass, where W is the biomass of a tree (kg of leaves, branches, stems or roots), a and b are constants and D is the DBH (cm). Other biomass components were sampled in different quadrats in three 1 hm² permanent research plots: shrubs (ten 25 m² quadrats), herbs (ten 4 m² quadrats), dead wood (the whole plot), large fallen branches (twenty-five 25 m² quadrats) and litterfall (twenty-five 1 m² quadrats). This method was used for estimating above- and below-ground biomass of live and dead plants (trees, seedlings, shrubs, herbs, woody lianas, epiphytes, coarse woody debris and litterfall).

Important findings Total biomass for the three plots was 370.163, 550.119 and 351.442 Mg·hm^-2, with an average of (423.908±109.702) Mg·hm^-2 (95% confidence interval). Living biomass made up 95.28% of the total biomass, with coarse woody debris and litterfall comprising the rest. Most living biomass (98.09%±0.60%) (Mean±SD, n=3) of the seasonal rain forest was concentrated in the tree layer. In the allocation of total biomass, stems accounted 68.33% and roots, branches and leaves made up 18.91%, 11.07% and 1.65%, respectively. The biomass allocation among different DBH classes was concentrated in the middle and largest classes, with large trees (D>70 cm) accounting for 43.67% ± 12.67%. The most important ten species, in terms of biomass, made up 63.43% of the tree layer. Leaf area index (LAI) of the tree layer for the three plots was 5.73, 7.35 and 6.08, with an average of 6.39. Estimated aboveground biomass in our study sites fell within the range of published values for tropical moist forests and was lower than that of Malaysian and Cameroon rain forests, but higher than some neotropical rain forests. In terms of total biomass, Xishuangbanna tropical seasonal rain forest is also higher than moist forest in Brazil.

Key words: biomass, biomass allocation, regression models, tropical seasonal rain forest, Xishuangbanna

吕晓涛, 唐建维, 何有才, 段文贵, 宋军平, 许海龙, 朱胜忠. 西双版纳热带季节雨林的生物量及其分配特征. 植物生态学报, 2007, 31(1): 11-22. DOI: 10.17521/cjpe.2007.0003

LÁ Xiao-Tao, TANG Jian-Wei, HE You-Cai, DUAN Wen-Gui, SONG Jun-Ping, XU Hai-Long, ZHU Sheng-Zhong. BIOMASS AND ITS ALLOCATION IN TROPICAL SEASONAL RAIN FOREST IN XISHUANGBANNA, SOUTHWEST CHINA. Chinese Journal of Plant Ecology, 2007, 31(1): 11-22. DOI: 10.17521/cjpe.2007.0003

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图/表 9

参考文献 37

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地点 Study site	样地号 Plot No.	纬度 Latitude	经度 Longitude	海拔 Altitude	坡向 Aspect	坡度 Slope	坡位 Position
勐仑 Menglun	Ⅰ	21°57' N	101°12' E	730 m	NW	15°~20°	中 Middle
勐腊 Mengla	Ⅱ	21°32' N	101°33' E	580 m	EN	25°~30°	下 Lower
曼养 Manyang	Ⅲ	21°27' N	101°36' E	640 m	NE	20°~25°	下 Lower

地点 Study site	样地号 Plot No.	纬度 Latitude	经度 Longitude	海拔 Altitude	坡向 Aspect	坡度 Slope	坡位 Position
勐仑 Menglun	Ⅰ	21°57' N	101°12' E	730 m	NW	15°~20°	中 Middle
勐腊 Mengla	Ⅱ	21°32' N	101°33' E	580 m	EN	25°~30°	下 Lower
曼养 Manyang	Ⅲ	21°27' N	101°36' E	640 m	NE	20°~25°	下 Lower

生活型 Life form	径级及样木数 DBH classes and sample no.	器官 Organ	回归模型 Regression models	相关系数 Regression coefficients (r)
乔木 Tree	2 cm≤D≤5 cm (N=46)	干 Stem	W_S=0.073 3D^{2.588 4}	0.896 0^***
		枝 Branch	W_B=0.013 5D^{2.515 8}	0.731 7^***
		叶 Leaf	W_L=0.039 4D^1.456	0.667 5^***
		根 Root	W_R=0.028D^2.399	0.826 6^***
		总 Total	W_T=0.147 1D^{2.439 9}	0.907 6^***
	5 cm<D≤20 cm (N=55)	干 Stem	W_S =0.108 6D^{2.316 9}	0.945 3^***
		枝 Branch	W_B =0.018 6D^{2.468 5}	0.861 9^***
		叶 Leaf	W_L =0.045 5D^{1.663 6}	0.767 5^***
		根 Root	W_R =0.024 2D^{2.420 5}	0.935 7^***
		总 Total	W_T =0.181 5D^{2.322 2}	0.959 1^***
	D>20 cm (N=22)	干 Stem	W_S=0.040 1D^{2.675 2}	0.966 3^***
		枝 Branch	W_B=0.082 9D^{2.039 5}	0.913 6^***
		叶 Leaf	W_L =0.097 9D^{1.358 4}	0.797 6^***
		根 Root	W_R =0.011 1D^{2.680 1}	0.968 6^***
		总 Total	W_T =0.093 2D^{2.560 2}	0.968 1^***
木质藤本 Woody liana	2 cm≤D≤14.5 cm (N=13)	总 Total	W_T=0.072 9(D²L)^{0.864 2}	0.942 3^***

生活型 Life form	径级及样木数 DBH classes and sample no.	器官 Organ	回归模型 Regression models	相关系数 Regression coefficients (r)
乔木 Tree	2 cm≤D≤5 cm (N=46)	干 Stem	W_S=0.073 3D^{2.588 4}	0.896 0^***
		枝 Branch	W_B=0.013 5D^{2.515 8}	0.731 7^***
		叶 Leaf	W_L=0.039 4D^1.456	0.667 5^***
		根 Root	W_R=0.028D^2.399	0.826 6^***
		总 Total	W_T=0.147 1D^{2.439 9}	0.907 6^***
	5 cm<D≤20 cm (N=55)	干 Stem	W_S =0.108 6D^{2.316 9}	0.945 3^***
		枝 Branch	W_B =0.018 6D^{2.468 5}	0.861 9^***
		叶 Leaf	W_L =0.045 5D^{1.663 6}	0.767 5^***
		根 Root	W_R =0.024 2D^{2.420 5}	0.935 7^***
		总 Total	W_T =0.181 5D^{2.322 2}	0.959 1^***
	D>20 cm (N=22)	干 Stem	W_S=0.040 1D^{2.675 2}	0.966 3^***
		枝 Branch	W_B=0.082 9D^{2.039 5}	0.913 6^***
		叶 Leaf	W_L =0.097 9D^{1.358 4}	0.797 6^***
		根 Root	W_R =0.011 1D^{2.680 1}	0.968 6^***
		总 Total	W_T =0.093 2D^{2.560 2}	0.968 1^***
木质藤本 Woody liana	2 cm≤D≤14.5 cm (N=13)	总 Total	W_T=0.072 9(D²L)^{0.864 2}	0.942 3^***

层次 Layer	样地 Plot	茎 Stem		枝 Branch		叶 Leaf		根 Root		总计 Total
层次 Layer	样地 Plot	生物量 Biomass	%	生物量 Biomass	%	生物量 Biomass	%	生物量 Biomass	%	生物量 Biomass	%
乔木层 Tree	I	238.396	68.50	37.733	10.84	4.418	1.27	67.493	19.39	348.040	100
	II	359.924	69.66	49.873	9.65	4.951	0.96	101.976	19.74	516.724	100
	III	219.764	67.61	38.379	11.81	4.655	1.43	62.239	19.15	325.037	100
	平均 Average	272.695 (76.115)	68.76	41.995 (6.830)	10.59	4.675 (0.267)	1.18	77.236 (21.586)	19.47	396.600 (104.664)	100
灌木层 Shrub	I	1.213	52.08	0.208	8.93	0.312	13.40	0.596	25.59	2.329	100
	II	0.609	53.33	0.106	9.28	0.155	13.57	0.272	23.82	1.142	100
	III	0.995	50.41	0.292	14.79	0.239	12.11	0.448	22.70	1.974	100
	平均 Average	0.939 (0.306)	51.73	0.202 (0.093)	11.13	0.235 (0.079)	12.97	0.439 (0.162)	24.17	1.815 (0.609)	100
草本层 Herb	I	0.534	35.82	-	-	0.328	22.0	0.629	42.19	1.491	100
	II	0.928	41.15	-	-	0.572	25.37	0.755	33.48	2.255	100
	III	0.455	18.56	-	-	0.628	25.61	1.369	55.83	2.452	100
	平均 Average	0.639 (0.253)	30.93	-	-	0.509 (0.160)	24.65	0.918 (0.396)	44.42	2.066 (0.508)	100
木质藤本 Liana	I	3.347	81.85	-	-	0.206	5.04	0.536	13.11	4.089	100
	II	2.469	87.71	-	-	0.152	5.40	0.194	6.89	2.815	100
	III	2.339	87.70	-	-	0.144	5.40	0.184	6.90	2.667	100
	平均 Average	2.718 (0.548)	85.21	-	-	0.167 (0.034)	5.25	0.305 (0.200)	9.55	3.190 (0.782)	100
附生植物 Epiphyte	I	0.115	50.66	-	-	0.065	28.63	0.047	20.71	0.227	100
	II	0.087	45.31	-	-	0.061	31.77	0.044	22.92	0.192	100
	III	0.139	45.57	-	-	0.098	32.13	0.068	22.30	0.305	100
	平均 Average	0.114 (0.026)	47.10	-	-	0.075 (0.020)	30.94	0.053 (0.013)	21.96	0.241 (0.058)	100
总计 Total	I	243.605	68.39	37.941	10.65	5.329	1.50	69.301	19.46	356.176	100
	II	364.017	69.59	49.979	9.55	5.891	1.13	103.241	19.73	523.128	100
	III	223.692	67.29	38.671	11.63	5.764	1.73	64.308	19.34	332.435	100
	平均 Average	277.105 (75.923)	68.61	42.197 (6.749)	10.45	5.661 (0.295)	1.40	78.950 (21.184)	19.55	403.913 (103.923)	100

西双版纳热带季节雨林的生物量及其分配特征

BIOMASS AND ITS ALLOCATION IN TROPICAL SEASONAL RAIN FOREST IN XISHUANGBANNA, SOUTHWEST CHINA

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径级 DBH classes (cm)	样地I Plot I		样地II Plot II		样地III Plot III		平均 Average
径级 DBH classes (cm)	个体数 No. of individual	生物量 Biomass	个体数 No. of individual	生物量 Biomass	个体数 No. of individual	生物量 Biomass	个体数 No. of individual	生物量 Biomass
2~5	1 562 (63.94)	4.309 (1.24)	1 127 (57.9)	3.131 (0.61)	1 163 (58.50)	2.87 (0.88)	1 284 (60.43)	3.44 (0.87)
5~10	488 (19.98)	8.156 (2.34)	396 (20.36)	7.03 (1.36)	359 (18.06)	6.433 (1.98)	414 (19.50)	7.21 (1.82)
10~20	205 (8.39)	17.424 (5.01)	218 (11.21)	20.106 (3.89)	227 (11.42)	21.876 (6.73)	217 (10.20)	19.802 (4.99)
20~30	91 (3.72)	30.528 (8.77)	88 (4.52)	29.610 (5.73)	117 (5.86)	38.342 (11.80)	99 (4.64)	32.827 (8.28)
30~40	42 (1.72)	33.579 (9.65)	39 (2.01)	33.092 (6.40)	63 (3.17)	45.950(14.14)	48 (2.26)	37.540 (9.47)
40~50	19 (0.78)	28.736 (8.26)	25 (1.29)	37.919 (7.34)	26 (1.31)	39.124 (12.04)	23 (1.10)	35.260 (8.89)
50~60	11 (0.45)	29.943 (8.60)	21 (1.08)	54.518 (10.55)	13 (0.65)	33.561 (10.32)	15 (0.71)	39.341 (9.92)
60~70	8 (0.33)	34.726 (9.98)	12 (0.62)	46.067 (8.92)	10 (0.50)	39.137 (12.04)	10 (0.47)	39.977 (10.08)
70~80	10 (0.41)	58.002 (16.67)	8 (0.41)	46.334 (8.97)	6 (0.30)	33.126 (10.19)	8 (0.38)	45.821 (11.55)
80~90	2 (0.08)	14.816 (4.26)	2 (0.10)	18.199 (3.52)	1 (0.05)	7.109 (2.19)	1.7 (0.08)	13.375 (3.37)
90~100	2 (0.08)	20.394 (5.86)	2 (0.10)	22.243 (4.30)	1 (0.05)	11.355 (3.49)	1.7 (0.08)	17.997 (4.54)
100~110	1 (0.04)	15.459 (4.44)	3 (0.15)	39.114 (7.57)	-	-	1.3 (0.06)	18.191 (4.59)
110~120	1 (0.04)	17.688 (5.08)	-	-	-	-	0.3 (0.02)	5.896 (1.49)
120~130	-	-	2 (0.10)	44.305 (8.57)	2 (0.10)	46.155 (14.20)	1.3 (0.06)	30.153 (7.60)
130~140	-	-	1 (0.05)	29.068 (5.62)	-	-	0.3 (0.02)	9.676 (2.44)
>140	1 (0.04)	34.280 (9.85)	1 (0.05)	86.029 (16.65)	-	-	0.7 (0.03)	40.089 (10.11)
总计Total	2 443 (100)	348.040 (100)	1 945 (100)	516.724 (100)	1 988 (100)	325.037 (100)	2 125 (100)	396.587 (100)

样地 Plot	树种 Species	植株个体 Individuals		生物量 Biomass
样地 Plot	树种 Species	个体密度 Density (No.·hm^-2)	占乔木层% % of the tree layer	Mg·hm^-2	占乔木层% % of the tree layer
I	绒毛番龙眼 Pometia tomentosa	70	2.87	87.355	25.10
	千果榄仁 Terminalia myriocarpa	1	0.04	34.280	9.85
	梭果玉蕊 Barringtonia macrostachya	90	3.68	20.307	5.83
	毛麻楝 Chukrasia tabularia var. velutina	10	0.41	17.711	5.09
	景洪猴欢喜 Sloanea cheliensis	11	0.45	13.278	3.81
	滇印杜英 Elaeocarpus varunua	7	0.28	13.186	3.79
	多花白头树 Garuga floribunda var. gamblai	9	0.37	12.899	3.71
	勐仑翅子树 Pterospermum menglunense	20	0.82	9.943	2.86
	思茅栲 Castanopsis ferox	3	0.12	9.671	2.78
	白颜树 Gironniera subaequalis	61	2.50	8.452	2.43
	总计Total	282	11.54	227.084	65.25
II	千果榄仁 Terminalia myriocarpa	6	0.31	154.254	29.85
	绒毛番龙眼 Pometia tomentosa	42	2.16	29.540	5.72
	皮孔葱臭木 Dysoxylum lenticellatum	31	1.59	27.229	5.27
	团花树 Anthocephalus chinensis	40	2.06	24.997	4.84
	多花白头树 Garuga floribunda var. gamblai	17	0.87	23.301	4.51
	龙果 Pouteria grandiflora	21	1.08	22.032	4.26
	黄棉木 Metadina trichotoma	8	0.41	19.395	3.75
	盆架树 Winchia calophylla	3	0.15	14.687	2.84
	琴叶风吹楠 Horsfieldia pandurifolia	22	1.13	13.900	2.69
	长叶榆 Ulmus lanceaefolia	7	0.36	13.260	2.57
	总计 Total	197	10.13	342.595	66.30
III	梭果玉蕊 Barringtonia macrostachya	166	8.35	44.555	13.71
	红果葱臭木 Dysoxylum binecteriferum	17	0.86	25.898	7.97
	浆果乌桕 Sapium baccatum	1	0.05	24.686	7.59
	粗壮琼楠 Beilschmiedia robusta	1	0.05	21.469	6.61
	白颜树 Gironniera subaequalis	63	3.17	17.693	5.44
	云南肉豆蔻 Myristica yunnanensis	165	8.30	13.720	4.22
	绒毛番龙眼 Pometia tomentosa	8	0.40	13.521	4.16
	多花白头树 Garuga floribunda var. gamblai	1	0.05	11.355	3.49
	云南黄杞 Engelhardtia spicata	4	0.20	10.199	3.14
	泰国黄叶树 Xanthophyllum siamensis	12	0.60	7.865	2.42
	总计 Total	438	22.03	190.961	58.75

样地 Plot	叶 Leaf		枝 Twig		花果 Fruit		杂屑 Other		总计Total
样地 Plot	生物量Biomass	%	生物量Biomass	%	生物量Biomass	%	生物量Biomass	%	生物量Biomass	%
I	1.848	62.39	0.809	27.31	0.284	9.59	0.020	0.68	2.962	100
II	1.103	36.41	1.819	60.05	0.072	2.38	0.035	1.16	3.028	100
III	1.110	50.68	0.938	42.83	0.113	5.16	0.029	1.32	2.190	100
平均 Average	1.354 (0.428)	49.65	1.189 (0.550)	43.60	0.156 (0.112)	5.72	0.028 (0.007)	1.03	2.727 (0.466)	100

生物量组分 Component	茎 Stem		枝 Branch		叶 Leaf		根 Root		总计 Total
生物量组分 Component	生物量 Biomass	%	生物量 Biomass	%	生物量 Biomass	%	生物量 Biomass	%	生物量 Biomass	%
活体生物量 Living biomass	277.105 (75.923)	68.61	42.197 (6.749)	10.45	5.661 (0.295)	1.40	78.950 (21.184)	19.55	403.913 (103.923)	100
凋落物 Litterfall	-	-	1.189 (0.550)	49.83	1.354 (0.428)	43.40	-	-	2.727^* (0.466)	100
粗死木质残体 CWD	12.547 (5.855)	72.66	3.522 (0.994)	20.39	-	-	1.199 (0.345)	6.94	17.268 (6.481)	100
总计 Total	289.652	68.33	46.908	11.07	7.015	1.65	80.149	18.91	423.908	100

森林类型 Forest type	地点 Location	面积 Area of plot (hm²)	最小胸径 Minimum DBH (cm)	乔木层地上生物量 AGB of tree layer	乔木层生物量 TB of tree layer	总生物量 TB	引用文献 References cited
低地湿性雨林LWR	圭亚那Guiana	22	10	309	-	-	Chave et al.,2001
低地湿润雨林LMR	巴拿马Panama	50	1	281	-	-	Chave et al.,2003
热带雨林TRF	巴西Brazil	392	-	264	-	372	Keller et al.,2001
热带湿润雨林TMF	喀麦隆Cameroon	10	15	312	-	-	Brown et al.,1989
热带湿润雨林TMF	斯里兰卡Sri Lanka	8	10	205	-	-
热带湿润雨林TMF	马来西亚Malaysia	0.5	10	391	-	-
热带湿润雨林TMF	柬埔寨Cambodia	-	10	295	-	-	Brown,1997
低地湿润雨林LMR	哥斯达黎加Costa Rica	0.3	1	234	-	-	Dewalt & Chave, 2004
山地雨林TMRF	海南Hainan	1	5	386	417	429	李意德等,1998 Li et al., 1998
山地雨林TMRF	海南Hainan	0.15	5.5	492	572	588	黄全等,1991 Huang et al., 1991
湿性季节雨林WSRF	西双版纳Xishuangbanna	0.25	5	536	683	693	郑征等,2000 Zheng et al., 2000
沟谷热带雨林RTRF	西双版纳Xishuangbanna	0.2	2	259	321	335	党承林等,1997 Dang et al., 1997
湿性季节雨林WSRF	西双版纳Xishuangbanna	1	5	283	353	361	冯志立等,1998 Feng et al., 1998
季节雨林TSRF	西双版纳Xishuangbanna	3	2	319	396	424	本研究 This study

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样地 Plot	枯立木 Standing dead trees		倒木 Fallen dead trees		大枝 Large dead branches		总计 Total
样地 Plot	生物量Biomass	%	生物量Biomass	%	生物量Biomass	%	生物量Biomass	%
I	5.548	50.32	2.148	19.48	3.329	30.20	11.025	100
II	5.222	21.79	14.143	59.02	4.598	19.19	23.963	100
III	8.128	48.33	6.050	35.98	2.638	15.69	16.817	100
平均 Average	6.299 (1.592)	36.48	7.447 (6.118)	43.13	3.522 (0.994)	20.40	17.268 (6.481)	100

样地 Plot	枯立木 Standing dead trees		倒木 Fallen dead trees		大枝 Large dead branches		总计 Total
样地 Plot	生物量Biomass	%	生物量Biomass	%	生物量Biomass	%	生物量Biomass	%
I	5.548	50.32	2.148	19.48	3.329	30.20	11.025	100
II	5.222	21.79	14.143	59.02	4.598	19.19	23.963	100
III	8.128	48.33	6.050	35.98	2.638	15.69	16.817	100
平均 Average	6.299 (1.592)	36.48	7.447 (6.118)	43.13	3.522 (0.994)	20.40	17.268 (6.481)	100