长白山温带森林不同演替阶段群落结构特征

doi:10.3773/j.issn.1005-264x.2009.06.009

摘要/Abstract

摘要：

原始阔叶红松林是长白山西部地区的地带性顶级植被类型, 经采伐干扰或火烧破坏后形成大面积次生林。参照CTFS (Centre for Tropical Forest Science)样地建设技术规范, 于2005~2007年, 在长白山地区典型次生杨桦林、次生针阔混交林和椴树红松林内各建立了5.2 hm²固定监测样地。调查并鉴定了样地内胸径大于1 cm的木本植物, 初步分析了森林监测样地的群落组成和种群结构, 并应用双相关函数g(r)分析了样地内5个优势树种的空间分布。结果表明: 次生杨桦林样地共监测木本植物32种, 20 949株活个体, 隶属于13科21属。次生针阔混交林样地共监测木本植物31个种, 14 725株活个体, 隶属于12科20属。椴树红松林样地共监测木本植物20个种, 12 062株活个体, 隶属于11科13属。次生杨桦林、次生针阔混交林及椴树红松林中胸径大于1 cm的木本植物胸高断面积之和分别为24.74、32.07和56.64 m²·hm^-2。紫椴(Tilia amurensis)是长白山针阔混交林带的重要组成树种, 其重要值、胸高断面积在3个森林监测样地内均居于前列。白桦(Betula platyphylla)、山杨(Populus daviana)重要值和胸高断面积在次生杨桦林内均处于优势地位, 而在椴树红松林内优势地位为红松(Pinus koraiensis)等顶级树种所取代。次生杨桦林和次生针阔混交林中, 红松、色木槭(Acer mono)、臭松(Abies nephrolepis)、鱼鳞松(Picea jezoensis)和紫椴的径级结构均呈倒J型分布; 而椴树红松林内, 红松和紫椴的径级结构则呈单峰分布, 色木槭、臭松和鱼鳞松呈倒J型分布。g(r)分析表明长白山森林监测样地内5个优势树种的空间格局以聚集分布为主, 聚集强度在同种个体周围(r≤4 m)达到最大, 随着距离增加, 聚集强度逐渐减小。次生林中树种空间格局的环境解释量较高, 而椴树红松林中环境因子对树种空间分布的解释能力较差。

关键词: 径级结构, 分布格局, 动态监测样地, 环境解释

Abstract:

Aims The Changbai Mountains had a large area of undisturbed temperate old-growth forest that was partly destroyed by large-scale industrial logging at the turn of the 20th century and replaced by secondary forests and plantations. Our objective was to further understand ecosystem development, community composition and structure, and spatial distribution of major tree populations.
Methods Three long-term 5.2 hm² forest plots were established in secondary Populus davidiana-Betula platyphylla forest (PBF), secondary conifer and broadleaved mixed forest (SCBMF) and Tilia amurensis-Pinus koraiensis mixed forest (TPMF) in the Changbai Mountains following the protocols of the Center for Tropical Forest Science forest dynamic plot. We tagged and recorded the species, diameter at breast height (DBH), height and crown diameter of all live and dead trees with DBH >1 cm. We determined the spatial distribution of five dominant canopy tree species by pair-correlation g(r) function and performed variation partitioning analysis to estimate the environmental effect on spatial distribution.
Important findings We recorded 20 949, 14 725 and 12 062 live individuals and basal areas of 24.74, 32.07 and 56.64 m²·hm^-2 in PBF, SCBMF and TPMF, respectively. Tilia amurensis, with the overall highest importance value and basal area, was the major dominant tree species in SCBMF, P. davidiana and B. platyphylla dominated PBF andP. koraiensis dominated TPMF. In the secondary forests BPF and SCBMF, the size-class distribution of Pinus koraiensis, Acer mono, Abies nephrolepis, Picea jezoensisandTilia amurensis showed the reverse J-shape form, whereas P. koraiensis and T. amurensis showed normal distributions in TPMF. Paired-correlation g(r) function showed that the main tree species had aggregated distributions, and the aggregation intensity decreased with the increasing distances. Environment variables had a strong effect on tree distribution, but the interpretation power of environment variables was very low in TPMF.

Key words: size-class distribution, spatial distribution pattern, dynamic monitoring forest plot, environmental interpretation

张春雨, 赵秀海, 赵亚洲. 长白山温带森林不同演替阶段群落结构特征. 植物生态学报, 2009, 33(6): 1090-1100. DOI: 10.3773/j.issn.1005-264x.2009.06.009

ZHANG Chun-Yu, ZHAO Xiu-Hai, ZHAO Ya-Zhou. COMMUNITY STRUCTURE IN DIFFERENT SUCCESSIONAL STAGES IN NORTH TEMPERATE FORESTS OF CHANGBAI MOUNTAINS, CHINA. Chinese Journal of Plant Ecology, 2009, 33(6): 1090-1100. DOI: 10.3773/j.issn.1005-264x.2009.06.009

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https://www.plant-ecology.com/CN/Y2009/V33/I6/1090

图/表 6

参考文献 27

[1]	Bonhomme R, Chartier P (1972). The interpretation and automatic measurement of hemispherical photographs to obtain sunlit folige area and gap frequency. Israel Journal of Agricultural Research, 22,53-61.
[2]	Borcard D, Legendre P, Drapeau P (1992). Partialling out the spatial component of ecological variation. Ecology, 73,1045-1055.
[3]	Borcard D, Legendre P (2002). All-scale spatial analysis of ecological data by means of principal coordinates of neighbour matrices. Ecological Modelling, 153,51-68.
[4]	Chen DK (陈大轲), Zhou XF (周晓峰), Zhu N (祝宁) (1994). Natural Secondary Forest: Structure, Function, Dynamic, and Management (天然次生林结构、功能动态与经营). Northeast Forestry University Press,Harbin.
[5]	China Soil Council (中国土壤学会) (1999). Soil Agricultural Chemical Analysis Procedure(土壤农业化学分析方法). Chinese Agricultural Science Press,Beijing. (in Chinese)
[6]	Center for Tropical Forest Science (2005). Methods for census of the 50 ha plot on Barro Colorado Island. http://www.ctfs.edu.
[7]	Condit R (1995). Research in large, long-term tropical forest plot. Trends in Ecological and Evolution, 10,18-22.
[8]	Condit R, Ashton PS, Baker P, Bunyavejchewin S, Gunatilleke S, Gunatilleke N, Hubbell SP, Foster RB, Itoh A, Lafrankie JV, Lee HS, Losos E, Manokaran N, Sukumar R, Yamakura T (2000). Spatial patterns in the distribution of tropical tree species. Science, 288,1414-1418. DOI URL PMID
[9]	Dray S, Legendre P, Peres-Neto PR (2006). Spatial modelling: a comprehensive framework for principal coordinate analysis of neighbour matrices (PCNM). Ecological Modelling, 196,483-493.
[10]	Engelbrecht BMJ, Comita LS, Condit R, Kursar TA, Tyree MT, Turner BL, Hubbell SP (2007). Drought sensitivity shapes species distribution patterns in tropical forests. Nature, 447,80-83. URL PMID
[11]	George LWP, Miller BP, Enright NJ (2006). A comparison of methods for the statistical analysis of spatial point patterns in plant ecology. Plant Ecology, 181,59-82.
[12]	Gerhardt K (1993). Tree seedling development in tropical dry abandoned pasture and secondary forest in Costa Rica. Journal of Vegetation Science, 4,95-102.
[13]	Gong GQ (宫贵权), Cheng JM (程积民), Mi XC (米湘成), Chen SW (陈声文), Fang T (方腾) (2007). Habitat associations of wood species in the Gutianshan subtropical broad-leaved evergreen forest. Science of Soil and Water Conservation (中国水土保持科学), 5,79-83. (in Chinese with English abstract)
[14]	Hao ZQ (郝占庆), Li BH (李步杭), Zhang J (张健), Wang XG (王绪高), Ye J (叶吉), Yao XL (姚晓琳) (2008). Broad-leaved Korean pine ( Pinus koraiensis) mixed forest plot in Changbaishan (CBS) of China: community composition and structure. Journal of Plant Ecology (Chinese Version)(植物生态学报), 32,238-250. (in Chinese with English abstract)
[15]	Hooper E, Condit R, Legendre P (2002). Responses of 20 native tree species to reforestation strategies for abandoned farmland in Panama. Ecological Applications, 12,1626-1641.
[16]	Lan GY (兰国玉), Hu YH (胡跃华), Cao M (曹敏), Zhu H (朱华), Wang H (王洪), Zhou SS (周仕顺), Deng XB (邓晓保), Cui JY (崔景云), Huang JG (黄建国), Liu LY (刘林云), Xu HL (许海龙), Song JP (宋军平), He YC (何有才) (2008). Establishment of Xishuangbanna tropical forest dynamics plot: species compositions and spatial distribution patterns. Journal of Plant Ecology (Chinese Version)(植物生态学报), 32,287-298. (in Chinese with English abstract)
[17]	Legendre P, Legendre L (2006). Numerical Ecology 2nd. Elsevier Science BV,Amsterdam.
[18]	Nepstad DC (1989). Forest Regrowth in Abandoned Pastures of Eastern Amazonia: Limitations to Tree Seedling Survival and Growth. PhD Dissertation, Yale University Press, New Haven.
[19]	Peres-Neto PR, Legendre P, Dray S, Borcard D (2006). Variation partitioning of species data matrices: estimation and comparison of fractions. Ecology, 87,2614-2625. DOI URL PMID
[20]	Rebertus AJ, Williamson GB, Moser EB (1989). Fire- induced changes in Quercus laevis spatial pattern in Florida sandhills. Journal of Ecology, 77,638-650.
[21]	Ripley BD (1981). Spatial Statistics. Wiley,New York.
[22]	Uhl C, Buschbacher R, Serrao EAS (1988). Abandoned pastures in eastern Amazonia. I. Patterns of plant succession. Journal of Ecology, 6,663-681.
[23]	Wiegand T, Moloney KA (2004). Rings, circles and null-models for point pattern analysis in ecology. Oikos, 104,209-229.
[24]	Ye WH (叶万辉), Cao HL (曹洪麟), Hang ZL (黄忠良), Lian JY (练琚愉), Wang ZG (王志高), Li L (李林), Wei SG (魏识广), Wang ZM (王章明) (2008). Community structure of a 20 hm ² lower subtropical evergreen broad-leaved forest plot in Dinghushan, China. Journal of Plant Ecology (Chinese Version)(植物生态学报), 32,274-286. (in Chinese with English abstract)
[25]	Zhang CY (张春雨), Zhao XH (赵秀海), Xia FC (夏富才) (2008). Spatial distribution of tree species and environmental interpretations in a secondary forest in Changbai Mountains. Scientia Silvae Sinicae (林业科学), 44,1-8. (in Chinese with English abstract)
[26]	Zhang CY, Gao LS, Zhao XH (2009). Spatial structures in a secondary forest in Changbai Mountains, Northeast China. Allgemeine Forst und Jagdzeitung(AFJZ), 314,45-55.
[27]	Zhu Y (祝燕), Zhao GF (赵谷风), Zhang LW (张俪文), Shen GC (沈国春), Mi XC (米湘成), Ren HB (任海保), Yu MJ (于明坚), Chen JH (陈建华), Chen SW (陈声文), Fang T (方腾), Ma KP (马克平) (2008). Community composition and structure of Gutianshan forest dynamic plot in a mid-subtropical evergreen broad-leaved forest, East China. Journal of Plant Ecology (Chinese Version)(植物生态学报), 32,262-273. (in Chinese with English abstract)

林型 Forest types	样地代码 Codes	经纬度 Longitude and latitude	平均海拔 Average altitude (m)	样地面积 Plot areas (hm²)	原始植被 Primary vegetation
次生杨桦林 Secondary Populus davidiana-Betula platyphylla forests	Plot 1	42o19.1667′ N 128o07.817′ E	899	5.2 (260 m×200 m)	阔叶红松林 Broad-leaved Pinus koraiensisforests
次生针阔混交林 Secondary conifer and broad-leaved mixed forests	Plot 2	42o20.907′ N 128o07.988′ E	748	5.2 (260 m×200 m)	阔叶红松林 Broad-leaved Pinus koraiensisforests
椴树红松林 Tilia amurensis-Pinus koraiensis mixed forests	Plot 3	42o13.684′ N 128o04.573′ E	1 042	5.2 (260 m×200 m)	阔叶红松林 Broad-leaved Pinus koraiensisforests

林型 Forest types	样地代码 Codes	经纬度 Longitude and latitude	平均海拔 Average altitude (m)	样地面积 Plot areas (hm²)	原始植被 Primary vegetation
次生杨桦林 Secondary Populus davidiana-Betula platyphylla forests	Plot 1	42o19.1667′ N 128o07.817′ E	899	5.2 (260 m×200 m)	阔叶红松林 Broad-leaved Pinus koraiensisforests
次生针阔混交林 Secondary conifer and broad-leaved mixed forests	Plot 2	42o20.907′ N 128o07.988′ E	748	5.2 (260 m×200 m)	阔叶红松林 Broad-leaved Pinus koraiensisforests
椴树红松林 Tilia amurensis-Pinus koraiensis mixed forests	Plot 3	42o13.684′ N 128o04.573′ E	1 042	5.2 (260 m×200 m)	阔叶红松林 Broad-leaved Pinus koraiensisforests

树种 Tree species	次生杨桦林 Secondary Populus davidiana-Betula platyphylla forests			次生针阔混交林 Secondary conifer and broadleaved mixed forests				椴树红松林 Tilia amurensis-Pinus koraiensis mixed forests
树种 Tree species	多度 Abun- dance	胸高断面积 Basal area (m²·hm^-2)	重要值Important value		多度 Abun-dance	胸高断面积Basal area (m²·hm^-2)	重要值Important value		多度 Abun- dance	胸高断面积 Basal area (m²·hm^-2)	重要值Important value
红松 Pinus koraiensis	439	0.99	3.79	709		2.06	6.35	911		28.55	24.62
紫椴 Tilia amurensis	2 268	3.58	10.66	1 103		6.30	11.64	796		15.55	16.74
糠椴 Tilia mandshurica	86	0.17	0.56	42		0.10	0.40	0		0	0
臭松 Abies nephrolepis	1 071	1.34	5.21	2616		2.65	11.28	3 847		4.05	13.02
蒙古栎 Quercus mongolica	727	1.49	4.98	237		2.41	5.06	1		<0.01	0.05
鱼鳞松 Picea jezoensis	508	1.46	4.73	192		0.83	2.40	214		0.44	3.71
水曲柳 Fraxinus mandshurica	773	3.22	7.54	285		2.58	4.65	0		0	0
色木槭 Acer mono	2 845	0.98	8.01	1 947		2.04	9.11	329		1.33	5.73
假色槭 Acer pseudo- sieboldianum	778	0.13	3.01	3 048		1.51	11.07	962		0.75	7.98
青楷槭 Acer tegmentosum	1 755	0.31	4.98	490		0.38	3.30	59		0.22	1.22
簇毛槭 Acer barbinerve	4 585	0.27	9.73	1 852		0.25	6.45	4 681		0.65	18.71
花楷槭 Acer ukurunduense	277	0.06	1.23	566		0.11	2.82	174		0.08	2.93
白牛槭 Acer mandshuricum	84	0.01	0.62	0		0	0.00	0		0	0
春榆 Ulmus japonica	754	1.03	4.41	167		0.17	1.77	0		0	0
怀槐 Maachia amurensis	175	0.18	1.33	260		0.80	3.30	0		0	0
黄檗 Phellodendron amurense	45	0.08	0.45	31		0.17	0.65	2		0.13	0.13
黄榆 Ulmus macrocarpa	0	0	0	17		0.05	0.29	2		<0.01	0.05
花楸 Sorbus pohuashanensis	63	0.05	0.59	25		0.04	0.46	1		<0.01	0.05
水榆 Sorbus alnifolia	83	0.20	0.89	26		0.08	0.44	1		<0.01	0.05
长白落叶松 Larix kaempferi	10	0.05	0.22	34		0.19	0.63	1		0.05	0.08
裂叶榆 Ulmus laciniata	128	0.10	1.11	33		0.03	0.47	3		0.01	0.10
山杨 Populus daviana	1 777	3.03	8.90	143		0.95	1.93	3		0.20	0.25
大青杨 Populus ussuriensis	64	0.57	1.18	45		2.86	4.31	71		4.52	4.42
香杨 Populus kareana	0	0	0	55		0.18	0.41	0		0	0
朝鲜柳 Salix koreensis	76	0.64	1.50	0		0	0.00	0		0	0
白桦 Betula platyphylla	733	3.69	7.98	643		5.23	8.91	2		0.04	0.04
枫桦 Betula costata	30	0.11	0.50	133		0.83	2.16	1		0.02	0.02
胡桃楸 Juglans mandshurica	42	0.06	0.48	0		0	0	0		0	0
色赤杨 Alnus tinctoria	233	0.76	2.12	0		0	0	0		0	0
大黄柳 Salix raddeana	19	<0.01	0.11	5		<0.01	0.08	0		0	0
毛山楂 Crataegus maximowiczii	0	0	0	6		<0.01	0.11	0		0	0
山丁子 Malus baccata	53	0.02	0.70	8		<0.01	0.16	0		0	0
稠李 Prunus padus	410	0.12	1.81	3		<0.01	0.07	0		0	0
鼠李 Rhamnus davurica	15	0.02	0.20	2		<0.01	0.04	0		0	0
黑樱桃 Cerasus maximowiczii	41	0.01	0.40	2		<0.01	0.04	0		0	0
红豆杉 Taxus cuspidata	0	0	0	0		0	0.00	1		<0.01	0.05
合计 Total	20 949	24.74		14 725		32.07		12 062		56.64

树种 Tree species	次生杨桦林 Secondary Populus davidiana-Betula platyphylla forests			次生针阔混交林 Secondary conifer and broadleaved mixed forests				椴树红松林 Tilia amurensis-Pinus koraiensis mixed forests
树种 Tree species	多度 Abun- dance	胸高断面积 Basal area (m²·hm^-2)	重要值Important value		多度 Abun-dance	胸高断面积Basal area (m²·hm^-2)	重要值Important value		多度 Abun- dance	胸高断面积 Basal area (m²·hm^-2)	重要值Important value
红松 Pinus koraiensis	439	0.99	3.79	709		2.06	6.35	911		28.55	24.62
紫椴 Tilia amurensis	2 268	3.58	10.66	1 103		6.30	11.64	796		15.55	16.74
糠椴 Tilia mandshurica	86	0.17	0.56	42		0.10	0.40	0		0	0
臭松 Abies nephrolepis	1 071	1.34	5.21	2616		2.65	11.28	3 847		4.05	13.02
蒙古栎 Quercus mongolica	727	1.49	4.98	237		2.41	5.06	1		<0.01	0.05
鱼鳞松 Picea jezoensis	508	1.46	4.73	192		0.83	2.40	214		0.44	3.71
水曲柳 Fraxinus mandshurica	773	3.22	7.54	285		2.58	4.65	0		0	0
色木槭 Acer mono	2 845	0.98	8.01	1 947		2.04	9.11	329		1.33	5.73
假色槭 Acer pseudo- sieboldianum	778	0.13	3.01	3 048		1.51	11.07	962		0.75	7.98
青楷槭 Acer tegmentosum	1 755	0.31	4.98	490		0.38	3.30	59		0.22	1.22
簇毛槭 Acer barbinerve	4 585	0.27	9.73	1 852		0.25	6.45	4 681		0.65	18.71
花楷槭 Acer ukurunduense	277	0.06	1.23	566		0.11	2.82	174		0.08	2.93
白牛槭 Acer mandshuricum	84	0.01	0.62	0		0	0.00	0		0	0
春榆 Ulmus japonica	754	1.03	4.41	167		0.17	1.77	0		0	0
怀槐 Maachia amurensis	175	0.18	1.33	260		0.80	3.30	0		0	0
黄檗 Phellodendron amurense	45	0.08	0.45	31		0.17	0.65	2		0.13	0.13
黄榆 Ulmus macrocarpa	0	0	0	17		0.05	0.29	2		<0.01	0.05
花楸 Sorbus pohuashanensis	63	0.05	0.59	25		0.04	0.46	1		<0.01	0.05
水榆 Sorbus alnifolia	83	0.20	0.89	26		0.08	0.44	1		<0.01	0.05
长白落叶松 Larix kaempferi	10	0.05	0.22	34		0.19	0.63	1		0.05	0.08
裂叶榆 Ulmus laciniata	128	0.10	1.11	33		0.03	0.47	3		0.01	0.10
山杨 Populus daviana	1 777	3.03	8.90	143		0.95	1.93	3		0.20	0.25
大青杨 Populus ussuriensis	64	0.57	1.18	45		2.86	4.31	71		4.52	4.42
香杨 Populus kareana	0	0	0	55		0.18	0.41	0		0	0
朝鲜柳 Salix koreensis	76	0.64	1.50	0		0	0.00	0		0	0
白桦 Betula platyphylla	733	3.69	7.98	643		5.23	8.91	2		0.04	0.04
枫桦 Betula costata	30	0.11	0.50	133		0.83	2.16	1		0.02	0.02
胡桃楸 Juglans mandshurica	42	0.06	0.48	0		0	0	0		0	0
色赤杨 Alnus tinctoria	233	0.76	2.12	0		0	0	0		0	0
大黄柳 Salix raddeana	19	<0.01	0.11	5		<0.01	0.08	0		0	0
毛山楂 Crataegus maximowiczii	0	0	0	6		<0.01	0.11	0		0	0
山丁子 Malus baccata	53	0.02	0.70	8		<0.01	0.16	0		0	0
稠李 Prunus padus	410	0.12	1.81	3		<0.01	0.07	0		0	0
鼠李 Rhamnus davurica	15	0.02	0.20	2		<0.01	0.04	0		0	0
黑樱桃 Cerasus maximowiczii	41	0.01	0.40	2		<0.01	0.04	0		0	0
红豆杉 Taxus cuspidata	0	0	0	0		0	0.00	1		<0.01	0.05
合计 Total	20 949	24.74		14 725		32.07		12 062		56.64

树种 Tree species	样地1 Plot 1			样地2 Plot 2			样地3 Plot 3
树种 Tree species	a	b	c	a	b	c	a	b	c
紫椴 Tilia amurensis	8.89	46.64	9.04	1.72	26.90	6.85	0	51.53	0
鱼鳞松 Picea jezoensis	1.63	45.36	0	27.45	68.10	29.98	0	62.71	0
色木槭 Acer mono	6.36	40.84	5.72	4.60	56.80	4.76	1.33	14.89	0
红松 Pinus koraiensis	3.09	66.04	4.26	5.35	38.35	4.99	1.34	44.29	0
臭松 Abies nephrolepis	10.18	78.22	12.09	12.75	50.74	13.29	0	64.71	0