Chin J Plant Ecol ›› 2020, Vol. 44 ›› Issue (3): 236-247.doi: 10.17521/cjpe.2019.0268

• Research Articles • Previous Articles     Next Articles

Association classification of a 30 hm2 dynamics plot in the monsoon broad-leaved evergreen forest in Pu’er, Yunnan, China

LI Shuai-Feng1,LANG Xue-Dong1,HUANG Xiao-Bo1,WANG Yan-Hong1,LIU Wan-De1,XU Chong-Hua2,SU Jian-Rong1,*()   

  1. 1Research Institute of Resource Insects, Chinese Academy of Forestry, The Pu'er Forest Ecosystem Research Station, National Forestry and Grassland Administration of China, Kunming 650224, China
    2Taiyanghe Provincinal Nature Reserve, Pu’er, Yunnan 665000, China
  • Received:2019-10-15 Accepted:2020-01-07 Online:2020-04-03 Published:2020-03-20
  • Contact: SU Jian-Rong
  • Supported by:
    Fundamental Research Funds of Chinese Academy of Forestry(CAFYBB2017ZX002-4);Forestry Industry Research Special Funds for Public Welfare Projects(201404211);Applied Basic Research Foundation of Yunnan Province(2019FB058)


Aims The monsoon broad-leaved evergreen forest is a typical zonal vegetation of south subtropical China, where forest dynamics plot is a study platform of mechanisms of biodiversity maintenance and community assembly. Our objectives were to obtain plant association types in a 30 hm2 forest dynamic monitoring plot in the Pu’er City, Yunnan Province.
Methods We identified typical association types by the methods of multivariate regression trees, importance value, principal component analysis and indicator species based on 750 subplots (20 m × 20 m), and analyzed the differences of topography factors and community characteristics among different plant associations.
Important findings The results showed that the specie richness of woody species (diameter at breast height (DBH) ≥ 1 cm) was 271, which belonged to 78 families and 178 genera in the forest dynamic plot. The community formation was Formation Castanopsis echidnocarpa. Forest was divided into four associations: (1) Lyonia ovalifolia + Aporosa villosa - Castanopsis echidnocarpa + Lithocarpus fenestratus Association, (2) Tapiscia yunnanensis + Lithocarpus grandifolius - Castanopsis echidnocarp + Schima wallichii Association, (3) Elaeocarpus sikkimensis + Polyspora chrysandra - Castanopsis echidnocarp + Schima wallichii Association, and (4) Betula alnoides + Alnus nepalensis - Castanopsis echidnocarp + Castanopsis calathiformis Association. There were many transitional overlaps in the distribution of associations, while indicator species was the main reference for distinguishing the types of associations. In addition, altitude and slope aspect had a great influence on the plant association types, while slope per se had a relatively weak impact on the association types.

Key words: vegetation classification, species composition, topography factor, multivariate regression tree, importance value, indicator species, monsoon broad-leaved evergreen forest

Fig. 1

Topography of the 30 hm2 forest dynamics plot of Pu’er. A, Contour map. B, Three dimensional topographic map."

Table 1

Stand characteristics of different life form in the 30 hm2 forest dynamics plot of Pu’er"

Species richness
Diameter at the
breast height (cm)
Height (m)
乔木 Tree 181 140 702 6.49 ± 7.25 6.98 ± 5.10
灌木 Shrub 39 5 271 2.85 ± 1.90 3.82 ± 1.77
藤本 Liana 51 8 399 3.84 ± 2.97 15.17 ± 6.17
合计 Total 271 154 372 6.22 ± 7.02 7.32 ± 5.46

Fig. 2

Species accumulation curves based on the cumulative subplot number in the 30 hm2 forest dynamics plot of Pu’er."

Fig. 3

Resubstitution and cross-validation relative error for the multivariate regression tree in the 30 hm2 forest dynamics plot of Pu’er. Min + 1SE, one standard error above the minimum cross-validated relative error; cp, complexity parameter; Inf, infinite."

Fig. 4

Multivariate regression tree and principal component analysis for association classification in the 30 hm2 forest dynamics plot of Pu’er. Red dots indicate Association 1 (Lyonia ovalifolia + Aporosa villosa - Castanopsis echidnocarpa + Lithocarpus fenestratus Association), green dots indicate Association 2 (Tapiscia yunnanensis + Lithocarpus grandifolius - Castanopsis echidnocarpa + Schima wallichii Association), dark blue dots indicate Association 3 (Elaeocarpus sikkimensis + Polyspora chrysandra - Castanopsis echidnocarpa + Schima wallichii Association), and light blue dots indicate Association 4 (Betula alnoides + Alnus nepalensis - Castanopsis echidnocarpa + Castanopsis calathiformis Association)."

Fig. 5

Four associations and venn diagram of species distribution at the scale of 20 m × 20 m in the 30 hm2 forest dynamics plot of Pu’er. Association 1, Lyonia ovalifolia + Aporosa villosa - Castanopsis echidnocarpa + Lithocarpus fenestratus Association; Association 2, Tapiscia yunnanensis + Lithocarpus grandifolius - Castanopsis echidnocarpa + Schima wallichii Association; Association 3, Elaeocarpus sikkimensis + Polyspora chrysandra - Castanopsis echidnocarpa + Schima wallichii Association; Association 4, Betula alnoides + Alnus nepalensis - Castanopsis echidnocarpa + Castanopsis calathiformis Association. Number in B represent species mumber."

Table 2

Order of four associations by importance value in the 30 hm2 forest dynamics plot of Pu’er"

群丛1 Association 1 群丛2 Association 2 群丛3 Association 3 群丛4 Association 4
物种 Species 重要值
物种 Species 重要值
物种 Species 重要值
物种 Species 重要值
1 短刺锥
Castanopsis echidnocarpa
27.80 短刺锥
Castanopsis echidnocarpa
10.28 短刺锥
Castanopsis echidnocarpa
25.75 短刺锥
Castanopsis echidnocarpa
2 泥柯
Lithocarpus fenestratus
9.99 西南木荷
Schima wallichii
6.96 西南木荷
Schima wallichii
9.55 枹丝锥
Castanopsis calathiformis
3 西南木荷
Schima wallichii
6.67 红梗润楠
Machilus rufipes
5.73 泥柯
Lithocarpus fenestratus
8.29 西南木荷
Schima wallichii
4 截果柯
Lithocarpus truncatus
5.88 云南瘿椒树
Tapiscia yunnanensis
5.38 枹丝锥
Castanopsis calathiformis
6.15 泥柯
Lithocarpus fenestratus
5 茶梨 Anneslea fragrans 4.23 泥柯
Lithocarpus fenestratus
4.73 红梗润楠
Machilus rufipes
5.87 红梗润楠
Machilus rufipes
6 西桦 Betula alnoides 4.06 齿叶黄杞
Engelhardtia serrata
4.01 截果柯
Lithocarpus truncatus
5.24 截果柯
Lithocarpus truncatus
7 红梗润楠
Machilus rufipes
4.03 耳叶柯
Lithocarpus grandifolius
3.56 红叶木姜子
Litsea rubescens
3.90 西桦 Betula alnoides 5.14
8 隐距越桔
Vaccinium exaristatum
3.95 厚皮香
Ternstroemia gymnanthera
2.97 茶梨
Anneslea fragrans
3.83 红叶木姜子
Litsea rubescens
9 红叶木姜子
Litsea rubescens
3.70 北酸脚杆
Medinilla septentrionalis
2.89 隐距越桔
Vaccinium exaristatum
1.98 云南瘿椒树
Tapiscia yunnanensis
10 枹丝锥
Castanopsis calathiformis
3.06 枹丝锥
Castanopsis calathiformis
2.89 深绿山龙眼
Helicia nilagirica
1.79 茶梨 Anneslea fragrans 2.56

Table 3

Order for four associations by species indicator value in the 30 hm2 forest dynamics plot of Pu’er"

Indicator value
Life form
Indicator value
Life form
群丛1 Association 1 毛杜茎山 Maesa permollis 0.208 T
红皮水锦树 Wendlandia tinctoria subsp. intermedia 0.529 T 构棘 Maclura cochinchinensis 0.206 L
隐距越桔 Vaccinium exaristatum 0.492 T 野柿 Diospyros kaki var. silvestris 0.195 T
深绿山龙眼 Helicia nilagirica 0.418 T 云南狗牙花 Ervatamia yunnanensis 0.172 T
巴豆藤 Craspedolobium unijugum 0.400 L 假山龙眼 Heliciopsis henryi 0.171 T
短刺锥 Castanopsis echidnocarpa 0.376 T 齿叶枇杷 Eriobotrya serrata 0.160 T
泥柯 Lithocarpus fenestratus 0.334 T 坚核桂樱 Laurocerasus jenkinsii 0.153 T
茶梨 Anneslea fragrans 0.320 T 粗梗胡椒 Piper macropodum 0.151 L
截果柯 Lithocarpus truncatus 0.308 T 苹果榕 Ficus oligodon 0.142 T
毛叶黄杞 Engelhardia spicata var. colebrookeana 0.264 T 珍珠伞 Ardisia maculosa 0.139 T
毛杨梅 Myrica esculenta 0.264 T 岭罗麦 Tarennoidea wallichii 0.138 T
珍珠花 Lyonia ovalifolia 0.243 T 刺篱木 Flacourtia indica 0.136 T
毛银柴 Aporosa villosa 0.194 T 台湾粗叶木 Lasianthus formosensis 0.135 T
猴耳环 Archidendron clypearia 0.188 T 假柿木姜子 Litsea monopetala 0.116 T
密花豆 Spatholobus suberectus 0.188 L 思茅锥 Castanopsis ferox 0.113 T
小果锥 Castanopsis fleuryi 0.168 T 分叉露兜 Pandanus urophyllus 0.112 T
密花树 Myrsine seguinii 0.147 T 焰序山龙眼 Helicia pyrrhobotrya 0.107 T
金叶子 Craibiodendron stellatum 0.113 T 棒果榕 Ficus subincisa 0.106 T
合果木 Michelia baillonii 0.107 T 越南安息香 Styrax tonkinensis 0.104 T
称杆树 Maesa ramentacea 0.077 T 四川冬青 Ilex szechwanensis 0.095 T
群丛2 Association 2 桫椤 Alsophila spinulosa 0.090 T
普文楠 Phoebe puwenensis 0.566 T 粗叶榕 Ficus hirta 0.088 S
阔叶蒲桃 Syzygium megacarpum 0.559 T 飞龙掌血 Toddalia asiatica 0.078 L
齿叶黄杞 Engelhardia serrata 0.514 T 望谟崖摩 Aglaia lawii 0.075 T
耳叶柯 Lithocarpus grandifolius 0.493 T 长叶柞木 Xylosma longifolia 0.074 T
北酸脚杆 Medinilla septentrionalis 0.466 T 砚壳花椒 Zanthoxylum dissitum 0.073 L
山香圆 Turpinia montana 0.462 T 鹧鸪花 Heynea trijuga 0.069 T
云南瘿椒树 Tapiscia yunnanensis 0.435 T 劲直刺桐 Erythrina stricta 0.065 T
艾胶树 Glochidion lanceolarium 0.423 T 鸡爪簕 Benkara sinensis 0.063 T
滇南木姜子 Litsea martabanica 0.393 T 剑叶木姜子 Litsea lancifolia 0.061 T
买麻藤 Gnetum montanum 0.358 L 尼泊尔水东哥 Saurauia napaulensis 0.060 T
钝叶桂 Cinnamomum bejolghota 0.356 T 腺萼木 Mycetia glandulosa 0.060 S
胭脂 Artocarpus tonkinensis 0.343 T 多蕊木 Tupidanthus calyptratus 0.059 T
独子藤 Celastrus monospermus 0.321 L 当归藤 Embelia parviflora 0.059 L
云南狗骨柴 Diplospora mollissima 0.320 T 毛叶合欢 Albizia mollis 0.052 T
纽子果 Ardisia virens 0.314 T 藤金合欢 Acacia concinna 0.051 L
李榄琼楠 Beilschmiedia linocieroides 0.302 T 红果樫木 Dysoxylum gotadhora 0.051 T
山杜英 Elaeocarpus sylvestris 0.295 T 玉兰叶木姜子 Litsea semecarpifolia 0.048 T
大叶斑鸠菊 Vernonia volkameriifolia 0.281 S 钩藤 Uncaria rhynchophylla 0.047 L
黑老虎 Kadsura coccinea 0.275 L 羊脆木 Pittosporum kerrii 0.047 T
山油柑 Acronychia pedunculata 0.242 T 榕叶掌叶树 Euaraliopsis ficifolia 0.042 S
异形南五味子 Kadsura heteroclita 0.240 L 小叶女贞 Ligustrum quihoui 0.042 T
瓦山锥 Castanopsis ceratacantha 0.222 T 中平树 Macaranga denticulata 0.041 T
山蕉 Mitrephora macclurei 0.216 T 微毛布惊 Vitex quinata var. puberula 0.040 T
越南石栎 Lithocarpus annamensis 0.214 T 歪叶榕 Ficus cyrtophylla 0.037 T
泰梭罗 Reevesia pubescens var. siamensis 0.032 T 群丛4 Association 4
森林榕 Ficus neriifolia 0.030 T 五瓣子楝树 Decaspermum parviflorum 0.408 T
矮小天仙果 Ficus erecta var. beecheyana 0.030 T 枹丝锥 Castanopsis calathiformis 0.320 T
南酸枣 Choerospondias axillaris 0.029 T 岗柃 Eurya groffii 0.280 T
爬树龙 Rhaphidophora decursiva 0.026 L 穗序鹅掌柴 Schefflera delavayi 0.260 T
鱼尾葵 Caryota maxima 0.024 T 鲫鱼胆 Maesa perlarius 0.253 T
扁担藤 Tetrastigma planicaule 0.024 L 三桠苦 Melicope pteleifolia 0.193 S
假苹婆 Sterculia lanceolata 0.023 T 圆锥悬钩子 Rubus paniculatus 0.192 L
浆果楝 Cipadessa baccifera 0.022 T 云南臀果木 Pygeum henryi 0.185 T
红枝崖爬藤 Tetrastigma erubescens 0.021 L 西桦 Betula alnoides 0.177 T
白花龙船花 Ixora henryi 0.020 T 黄心树 Machilus gamblei 0.170 T
多体蕊黄檀 Dalbergia polyadelpha 0.020 T 奶桑 Morus macroura 0.148 T
割舌树 Walsura robusta 0.020 T 喜马拉雅崖爬藤 Tetrastigma rumicispermum 0.146 L
水麻 Debregeasia orientalis 0.020 S 檀梨 Pyrularia edulis 0.115 T
泰国黄叶树 Xanthophyllum flavescens 0.020 T 红叶藤 Rourea minor 0.089 L
瓦理棕 Wallichia gracilis 0.020 T 云南黄杞 Engelhardia spicata 0.086 T
牛枓吴萸 Tetradium trichotomum 0.018 T 玉叶金花 Mussaenda pubescens 0.083 L
樱叶杜英 Elaeocarpus prunifolioides 0.018 T 托叶黄檀 Dalbergia stipulacea 0.078 L
细花秀丽火把花 0.017 L 大血藤 Sargentodoxa cuneata 0.077 L
Colquhounia elegans var. tenuiflora 无腺吴萸 Tetradium fraxinifolium 0.074 T
群丛3 Association 3 尼泊尔桤木 Alnus nepalensis 0.070 T
红叶木姜子 Litsea rubescens 0.310 T 高盆樱桃 Cerasus cerasoides 0.065 T
西南木荷 Schima wallichii 0.301 T 小萼瓜馥木 Fissistigma polyanthoides 0.057 L
红花木犀榄 Olea rosea 0.218 T 七小叶崖爬藤 Tetrastigma delavayi 0.038 L
腺叶木犀榄 Olea paniculata 0.198 T 一担柴 Colona floribunda 0.025 T
猪肚木 Canthium horridum 0.187 T 楹树 Albizia chinensis 0.022 T
大果杜英 Elaeocarpus sikkimensis 0.165 T 白瑞香 Daphne papyracea 0.020 S
黄药大头茶 Polyspora chrysandra 0.104 T 爪哇黄杞 Engelhardia spicata var. aceriflora 0.017 T
青藤仔 Jasminum nervosum 0.017 L

Table 4

Variance among four associations in the 30 hm2 forest dynamics plot of Pu’er (mean ± SE)"

类型 Type 面积
Area (hm2)
Species richness
Stand density
Diameter at breast height (cm)
Altitude (m)
Slope (°)
Slope aspect (°)
群丛1 Association 1 16.20 195 5 381.79 6.29 ± 0.71b 1 544.24 ± 14.92a 18.7 ± 5.07b 196.01 ± 88.33c
群丛2 Association 2 4.00 214 3 851.50 6.91 ± 1.62a 1 477.81 ± 5.32c 20.22 ± 6.84a 268.73 ± 70.58b
群丛3 Association 3 5.04 186 7 847.00 6.22 ± 0.79b 1 506.53 ± 9.64b 20.17 ± 3.93a 163.88 ± 46.86d
群丛4 Association 4 4.76 172 2 370.00 6.2 ± 0.89b 1 506.22 ± 10.39b 18.42 ± 5.58b 293.76 ± 43.71a

Table 5

Comparison of forest dynamic plots between Pu’er and adjacent districts"

类型 Type 森林动态监测样地 Forest dynamic plot
普洱 Pu’er 西双版纳 Xishuangbanna 哀牢山 Ailaoshan
建成时间 Establishment time (year) 2019 2007 2014
植被类型 Vegetation types 季风常绿阔叶林
Monsoon broad-leaved evergreen
Tropical forest
Mid-moutain moist evergreen
broad-leaved forest
面积 Area (hm2) 30 20 20
物种丰富度 Species richness 271 468 104
海拔范围 Altitude range (m) 1 467-1 587 709-869 2 472-2 628
优势物种 Dominant species 短刺锥、西南木荷、枹丝锥、泥柯、截果柯、红梗润楠、茶梨、西桦、耳叶柯
Castanopsis echidnocarpa, Schima wallichii, Castanopsis calathiformis, Lithocarpus fenestratus, Lithocarpus truncates, Machilus rufipes, Anneslea fragrans, Betula alnoides, Lithocarpus grandifolius
Pittosporopsis kerrii, Parashorea chinensis, Garcinia cowa, Castanopsis echidnocarpa, Orophea laui, Sloanea tomentosa, Baccaurea ramiflora, Knema tenuinervia, Pometia pinnata
Camellia forrestii, Vaccinium duclouxii, Symplocos ramosissima, Lithocarpus hancei, Castanopsis wattii, Eriobotrya bengalensis, Lithocarpus xylocarpus, Symplocos sumuntia, Machilus yunnanensis
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