基于浙江天童20 hm2常绿阔叶林动态监测样地的群丛划分

doi:10.17521/cjpe.2017.0263

摘要/Abstract

摘要：

为获取能够代表浙江天童山的森林植被典型群丛类型, 同时也为植被分类中如何发现过渡类型和确定典型类型提供参考, 该研究利用天童20 hm ²森林大样地资料, 运用双向指示种分析(TWINSPAN)与除趋势对应分析(DCA), 剔除过渡群落, 进行群丛划分。结果表明, 去除过渡地段后更利于研究区域典型群丛类型的确定。大样地的植被类型可划分为宜昌荚蒾-厚皮香/港柯+云山青冈群丛(Viburnum erosum-Ternstroemia gymnanthera/Lithocarpus harlandii + Cyclobalanopsis sessilifolia Ass.); 虎皮楠-柯/木荷+米槠群丛(Daphniphyllum oldhami-Lithocarpus glaber/Schima superba + Castanopsis carlesii Ass.); 红毒茴-紫楠/南酸枣+薄叶润楠群丛(Illicium lanceolatum-Phoebe sheareri/Choerospondias axillaries + Machilus leptophylla Ass.)。DCA排序同时能反映各群丛类型分布与环境的相关关系, 结果显示, 海拔和凹凸度对群丛分布有较大影响, 坡度和坡向对群丛分布影响较小。

关键词: 植被分类, 过渡群落, 种类组成, 双向指示种分析, 除趋势对应分析, 生境, 天童

Abstract:

Aims Our objectives were to obtain typical plant association types in a 20 hm ² forest dynamic monitoring plot in Mt. Tiantong, and provide methodological references for determining transitional types and identifying typical types in vegetation classification.

Methods We used two-way indicator species analysis (TWINSPAN) and detrended correspondence analysis (DCA) to remove transitional community and identify typical association types. DCA method was also used to analyze the relationship between the distributions of different plant associations and environmental variables.

Important findings The results showed that the classification of typical plant associations was greatly improved by the removal of transitional communities. The forest was divided into three associations: 1) Viburnum erosum- Ternstroemia gymnanthera/Lithocarpus harlandii + Cyclobalanopsis sessilifolia Association, 2) Daphniphyllum oldhami-Lithocarpus glaber/Schima superba + Castanopsis carlesii Association, and 3) Illicium lanceolatum- Phoebe sheareri/Choerospondias axillaries + Machilus leptophylla Association. The results of DCA revealed the importance of environmental variables (elevation, convexity, slope, and aspect) on spatial distribution of plant associations. Elevation and convexity had significant effects on the distribution of associations, while slope and aspect had weak effects. The results confirmed the high accuracy of the association classification we used.

Key words: vegetation classification, transitional community, floristic composition, two-way indicator species analysis, detrended correspondence analysis, habitat, Mt. Tiantong

孙小伟, 杨庆松, 刘何铭, 王希华. 基于浙江天童20 hm²常绿阔叶林动态监测样地的群丛划分. 植物生态学报, 2018, 42(5): 550-561. DOI: 10.17521/cjpe.2017.0263

SUN Xiao-Wei, YANG Qing-Song, LIU He-Ming, WANG Xi-Hua. Classification of plant associations based on a 20 hm² dynamics plot of evergreen broad- leaved forest in Mt. Tiantong, Zhejiang, China. Chinese Journal of Plant Ecology, 2018, 42(5): 550-561. DOI: 10.17521/cjpe.2017.0263

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2017.0263

https://www.plant-ecology.com/CN/Y2018/V42/I5/550

图/表 14

图1 浙江省天童20 hm2森林动态样地所在位置及三维地形图。

Fig. 1 The location and topography of the 20 hm2 forest dynamics plot in Mt. Tiantong, Zhejiang Province.

图2 物种分布不均匀的样方(A, 样方编号83, B, 样方编号324)。圆点表示样方中的植物个体。

Fig. 2 Quadrats with uneven distribution of species (A, stands for quadrat 83; B, stands for quadrat 324). Dots represent the plant individuals in each quadrat.

图3 天童20 hm2样地中用于分类的20 m × 20 m的样方分布图(198个样方, 等高线间距为10 m)。灰色阴影部分为林窗, 含有阴影圆圈的样方为用于分类的样方。

Fig. 3 Map of quadrats for classification at the scale of 20 m × 20 m in Tiantong 20 hm2 plot (198 quadrats in total) (The contour lines with 10 m intervals were shown). Gray polygons represent forest gaps, and shadow circles stand for the quadrats used for classification.

图4 天童20 hm2样地内用于分类的198个样方的TWINSPAN分类结果树状示意图。N表示样方总数, 方框里的物种表示各分类水平的正负指示种, 两者用“/”区分。CARVIM, 雷公鹅耳枥; CASCAR, 米槠; DISMYR, 杨梅叶蚊母树; EURLOQ, 细枝柃; EURRUB, 窄基红褐柃; LITELO, 黄丹木姜子; MACLEP, 薄叶润楠; MYRRUB, 杨梅; PHOSHE, 紫楠; SCHSUP, 木荷; SYMANO, 薄叶山矾。

Fig. 4 The result of TWINSPAN classification for 198 quadrats in Tiantong 20 hm2 plot. N represents total number of quadrats. Species in the box are indicator species, and a negative group and a positive group are included at each level of classification, they are separated by “/”. CARVIM, Carpinus viminea; CASCAR, Castanopsis carlesii; DISMYR, Distylium myricoides; EURLOQ, Eurya loquaiana; EURRUB, Eurya rubiginosa var. attenuata; LITELO, Litsea elongata; MACLEP, Machilus leptophylla; MYRRUB, Myrica rubra; PHOSHE, Phoebe sheareri; SCHSUP, Schima superba; SYMANO, Symplocos anomala.

图5 天童20 hm2样地5个类群的20 m × 20 m的样方分布图。A, B, C, D, E分别表示表示类群A-E包含的样方在样地中的分布。

Fig. 5 Distributions of five clusters at the scale of 20 m × 20 m in Tiantong 20 hm2 plot. A, B, C, D, E represent the distributions of the quadrats from clusters A-E, respectively.

图6 天童20 hm2样地内用于分类的198个样方的DCA排序图。阿拉伯数字表示样方编号。A, 类群A; B, 类群B; C, 类群C; D, 类群D; E, 类群E。

Fig. 6 DCA ordination diagram for the classification of 198 quadrats in Tiantong 20 hm2 plot with the five clusters. The numbers in the figure are the identification code for each quadrat. A, B, C, D, E represent the distributions of the quadrats in clusters A-E, respectively.

图7 去除过渡群落后剩余78个样方的TWINSPAN分类结果树状示意图。N表示样方总数, EURLOQ, 细枝柃; MACLEP, 薄叶润楠。

Fig. 7 The results of TWINSPAN classification for 78 quadrats after removing the transitional quadrats. N represents total number of quadrats. EURLOQ, Eurya loquaiana; MACLEP, Machilus leptophylla.

图8 天童20 hm2样地3种群落类型的20 m × 20 m的样方分布图。, , 分别表示群落1、群落2、群落3中包含的样方在样地中的分布。

Fig. 8 Distributions of three community types at the scale of 20 m × 20 m in Tiantong 20 hm2 plot. ,, and represent the distributions of the communities 1-3, respectively.

图9 去除过渡群落后剩余78个样方的DCA排序图。阿拉伯数字表示样方编号。a, 群落1; b, 群落2; c, 群落3。

Fig. 9 DCA ordination diagram of 78 quadrats after removing the transitional quadrats. The numbers in the figure are the identification code for each quadrat. a, community 1; b, community 2; c, community 3.

图10 3种群落类型的轮廓宽度值图。每一条水平线的长度代表一个样方的轮廓宽度值。n, 样方总数; j, 各类型编号; nj, 各群落内的样方总数; avei∈cj si, 各群落内的平均轮廓宽度值。

Fig. 10 The silhouette width of three community types. The length of each horizontal line represents the silhouette width of one quadrat. n, total number of quadrats; j, cluster number; nj, total number of quadrats in the community j; avei∈cj si, the average silhouette width in the community j.

表1 天童20 hm2样地3种群落类型内物种的重要值排序(前5)

Table 1 The order of species important value in three community types in Tiantong 20 hm2 plot (top 5)

物种 Species	重要值 Important value (%)	生长型 Growth form
群落1 Community 1
港柯 Lithocarpus harlandii	12.20	乔木 Tree
杨梅叶蚊母树 Distylium myricoides	10.97	小乔木 Small tree
云山青冈 Cyclobalanopsis sessilifolia	9.27	乔木 Tree
木荷 Schima superba	6.10	乔木 Tree
马银花 Rhododendron ovatum	6.06	小乔木 Small tree
群落2 Community 2
细枝柃 Eurya loquaiana	13.38	小乔木 Small tree
木荷 Schima superba	12.46	乔木 Tree
杨梅叶蚊母树 Distylium myricoides	12.12	小乔木 Small tree
米槠 Castanopsis carlesii	10.04	乔木 Tree
栲 Castanopsis fargesii	6.01	乔木Tree
群落3 Community 3
黄丹木姜子 Litsea elongata	21.46	小乔木 Small tree
薄叶润楠 Machilus leptophylla	12.06	乔木 Tree
南酸枣 Choerospondias axillaris	10.48	乔木 Tree
细枝柃 Eurya loquaiana	9.34	小乔木 Small tree
云山青冈 Cyclobalanopsis sessilifolia	5.90	乔木 Tree

表2 3种群落类型内物种指示值(大于0.5)排序

Table 2 The order of species indicator value (>0.5) in three community types in Tiantong 20 hm2 plot

物种 Species	特异性Specificity	确限度Fidelity	指示值Indicator value	p
群落1 Community 1
杜鹃 Rhododendron simsii	0.9659	0.7037	0.824	0.001^***
宜昌荚蒾 Viburnum erosum	0.9298	0.5556	0.719	0.001^***
厚皮香 Ternstroemia gymnanthera	0.9643	0.4444	0.655	0.001^***
迎春樱桃 Cerasus discoidea	1.0000	0.3704	0.609	0.001^***
山鸡椒 Litsea cubeba	0.9070	0.4074	0.608	0.002^**
青冈 Cyclobalanopsis glauca	0.8421	0.3704	0.558	0.003^**
大青 Clerodendrum cyrtophyllum	0.8750	0.3333	0.540	0.007^**
群落2 Community 2
虎皮楠 Daphniphyllum oldhami	0.9894	0.6957	0.830	0.001^***
柯 Lithocarpus glaber	0.9792	0.6087	0.772	0.001^***
杨梅 Myrica rubra	0.8667	0.6087	0.726	0.001^***
总状山矾 Symplocos botryantha	0.9375	0.4783	0.670	0.001^***
刺毛越桔 Vaccinium trichocladum	0.8889	0.4783	0.652	0.001^***
铁冬青 Ilex rotunda	0.5833	0.5217	0.552	0.045^*
群落3 Community 3
薄叶润楠 Machilus leptophylla	0.9931	1.0000	0.997	0.001^***
红毒茴 Illicium lanceolatum	0.9695	0.8214	0.892	0.001^***
紫楠 Phoebe sheareri	1.0000	0.5714	0.756	0.001^***
黄牛奶树 Symplocos cochinchinensis var. laurina	0.8482	0.6429	0.738	0.001^***
西川朴 Celtis vandervoetiana	1.0000	0.3214	0.567	0.001^***
锐角枫 Acer acutum	1.0000	0.2500	0.500	0.002^**
胡桃楸 Juglans mandshurica	1.0000	0.2500	0.500	0.001^***

图11 78个样方的DCA排序轴与环境因子的相关性图。阿拉伯数字表示样方编号。a, 宜昌荚蒾-厚皮香/港柯+云山青冈群丛; b, 虎皮楠-柯/木荷+米槠群丛; c, 红毒茴-紫楠/南酸枣+薄叶润楠群丛。

Fig. 11 DCA ordination diagram of 78 quadrats with environmental variables. The numbers in the figure are the identification code for each quadrat. a, Viburnum erosum-Ternstroemia gymnanthera/Lithocarpus harlandii + Cyclobalanopsis sessilifolia Association; b, Daphniphyllum oldhami-Lithocarpus glaber/Schima superba + Castanopsis carlesii Association; c, Illicium lanceolatum-Phoebe sheareri/Choerospondias axillaris + Machilus leptophylla Association.

表3 5个类群内物种指示值排序

Table 3 The order of species indicator value in five clusters in Tiantong 20 hm2 plot

物种 Species	特异性Specificity	确限度Fidelity		指示值Indicator value		p
类群A Cluster A
杜鹃 Rhododendron simsii	0.8788	0.6667		0.765		0.001^***
宜昌荚蒾 Viburnum erosum	0.8281	0.5000		0.643		0.001^***
厚皮香 Ternstroemia gymnanthera	0.8056	0.5000		0.635		0.001^***
山鸡椒 Litsea cubeba	0.5362	0.4000		0.463		0.001^***
迎春樱桃 Cerasus discoidea	0.6071	0.3333		0.450		0.001^***
南烛 Vaccinium bracteatum	0.5143	0.3333		0.414		0.003^**
豆腐柴 Premna microphylla	0.6000	0.1333		0.283		0.045^*
野鸦椿 Euscaphis japonica	0.7500	0.1000		0.274		0.026^*
木蜡树 Toxicodendron sylvestre	1.0000	0.0667		0.258		0.029^*
类群B Cluster B
柯 Lithocarpus glaber	0.7833	0.6087		0.691		0.001^***
杨梅 Myrica rubra	0.6047	0.6087		0.607		0.001^***
刺毛越桔 Vaccinium trichocladum	0.5333	0.4783		0.505		0.001^***
总状山矾 Symplocos botryantha	0.4688	0.4783		0.473		0.001^***
类群E Cluster E
紫楠 Phoebe sheareri	0.9000	0.5806		0.723		0.001^***
胡桃楸 Juglans mandshurica	0.9167	0.2581		0.486		0.001^***
锐角槭 Acer acutum	0.8333	0.2258		0.434		0.001^***
中华卫矛 Euonymus nitidus	0.7000	0.1613		0.336		0.006^**
枳椇 Hovenia acerba	0.8750	0.1290		0.336		0.002^**
类群B + C Cluster B + C
虎皮楠 Daphniphyllum oldhami	0.8685	0.5889		0.715		0.001^***
皱柄冬青 Ilex kengii	0.7812	0.1889		0.384		0.013^*
类群D + E Cluster D + E
薄叶润楠 Machilus leptophylla	0.9809	0.7436		0.854		0.001^***
红毒茴 Illicium lanceolatum	0.9031	0.6539		0.768		0.001^***
黄牛奶树 Symplocos cochinchinensis var. laurina	0.8273	0.5513		0.675		0.001^***
西川朴 Celtis vandervoetiana	0.9730	0.3077		0.547		0.001^***
无患子 Sapindus saponaria	0.8529	0.2564		0.468		0.001^***
糙叶树 Aphananthe aspera	0.8846	0.2308		0.452		0.001^***
紫弹树 Celtis biondii	0.8889	0.1923		0.413		0.003^**
杭州榆 Ulmus changii	0.9853	0.1410		0.373		0.007^**
青钱柳 Cyclocarya paliurus	0.8667	0.1282		0.333		0.009^**
大叶旱樱 Cerasus subhirtella	0.8750		0.0897		0.280	0.04^*
黐花 Mussaenda esquirolii	1.0000		0.0769		0.277	0.036^*

表3 5个类群内物种指示值排序

Table 3 The order of species indicator value in five clusters in Tiantong 20 hm2 plot

物种 Species	特异性Specificity	确限度Fidelity		指示值Indicator value		p
类群A Cluster A
杜鹃 Rhododendron simsii	0.8788	0.6667		0.765		0.001^***
宜昌荚蒾 Viburnum erosum	0.8281	0.5000		0.643		0.001^***
厚皮香 Ternstroemia gymnanthera	0.8056	0.5000		0.635		0.001^***
山鸡椒 Litsea cubeba	0.5362	0.4000		0.463		0.001^***
迎春樱桃 Cerasus discoidea	0.6071	0.3333		0.450		0.001^***
南烛 Vaccinium bracteatum	0.5143	0.3333		0.414		0.003^**
豆腐柴 Premna microphylla	0.6000	0.1333		0.283		0.045^*
野鸦椿 Euscaphis japonica	0.7500	0.1000		0.274		0.026^*
木蜡树 Toxicodendron sylvestre	1.0000	0.0667		0.258		0.029^*
类群B Cluster B
柯 Lithocarpus glaber	0.7833	0.6087		0.691		0.001^***
杨梅 Myrica rubra	0.6047	0.6087		0.607		0.001^***
刺毛越桔 Vaccinium trichocladum	0.5333	0.4783		0.505		0.001^***
总状山矾 Symplocos botryantha	0.4688	0.4783		0.473		0.001^***
类群E Cluster E
紫楠 Phoebe sheareri	0.9000	0.5806		0.723		0.001^***
胡桃楸 Juglans mandshurica	0.9167	0.2581		0.486		0.001^***
锐角槭 Acer acutum	0.8333	0.2258		0.434		0.001^***
中华卫矛 Euonymus nitidus	0.7000	0.1613		0.336		0.006^**
枳椇 Hovenia acerba	0.8750	0.1290		0.336		0.002^**
类群B + C Cluster B + C
虎皮楠 Daphniphyllum oldhami	0.8685	0.5889		0.715		0.001^***
皱柄冬青 Ilex kengii	0.7812	0.1889		0.384		0.013^*
类群D + E Cluster D + E
薄叶润楠 Machilus leptophylla	0.9809	0.7436		0.854		0.001^***
红毒茴 Illicium lanceolatum	0.9031	0.6539		0.768		0.001^***
黄牛奶树 Symplocos cochinchinensis var. laurina	0.8273	0.5513		0.675		0.001^***
西川朴 Celtis vandervoetiana	0.9730	0.3077		0.547		0.001^***
无患子 Sapindus saponaria	0.8529	0.2564		0.468		0.001^***
糙叶树 Aphananthe aspera	0.8846	0.2308		0.452		0.001^***
紫弹树 Celtis biondii	0.8889	0.1923		0.413		0.003^**
杭州榆 Ulmus changii	0.9853	0.1410		0.373		0.007^**
青钱柳 Cyclocarya paliurus	0.8667	0.1282		0.333		0.009^**
大叶旱樱 Cerasus subhirtella	0.8750		0.0897		0.280	0.04^*
黐花 Mussaenda esquirolii	1.0000		0.0769		0.277	0.036^*

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基于浙江天童20 hm²常绿阔叶林动态监测样地的群丛划分

Classification of plant associations based on a 20 hm² dynamics plot of evergreen broad- leaved forest in Mt. Tiantong, Zhejiang, China

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