生物多样性 ›› 2016, Vol. 24 ›› Issue (12): 1353-1363. DOI: 10.17520/biods.2016095
收稿日期:
2016-04-01
接受日期:
2016-10-19
出版日期:
2016-12-20
发布日期:
2017-01-10
通讯作者:
骆争荣
基金资助:
Yunxia Huang, Xuan Xu, Lixiang Zhang, Yue Song, Zhengrong Luo*()
Received:
2016-04-01
Accepted:
2016-10-19
Online:
2016-12-20
Published:
2017-01-10
Contact:
Luo Zhengrong
摘要:
森林群落的动态受多方面因素的影响。林下的草本植物和木本植物幼苗幼树对于森林的更新及动态具有重要作用。为了解百山祖常绿阔叶林群落下层植物物种组成和分布的动态变化, 我们分析了2003-2013年间百山祖5 ha森林动态样地灌草层物种(包括DBH < 1 cm的木本植物和所有草本植物)在物种重要值、α多样性以及物种分布等方面的变化。结果表明: (1) 10年间, 群落灌草层植物个体数减少了28.7%, 其中草质藤本植物个体数下降比例最大(76.9%), 而乔木的幼苗和幼树下降比例最小(14.4%); (2)整个灌草层的物种数明显下降, 同时乔木的幼苗和幼树及直立灌木的均匀度均下降, 而草本植物均匀度有所上升; (3)除短尾柯(Lithocarpus brevicaudatus)、叶萼山矾(Symplocos phyllocalyx)、显脉野木瓜(Stauntonia conspicua)和福建悬钩子(Rubus fujianensis)等植物重要值逆势上扬外, 直立灌木物种重要值10年间的变化值与其在2003年时的重要值呈显著正相关, 而其余各类生活型植物都显示, 初始重要值越高的物种在10年后下降幅度越大; (4)物种分布与生境类型的相关性变化复杂, 除与山谷生境正相关的物种数量增加外, 研究期间与生境存在显著相关性的物种数都有所减少; (5) 10年来仅陡坡处物种多样性指数没有显著下降。总体来说, 百山祖常绿阔叶林灌草层群落仍未达到顶极状态, 2003-2013年间正缓慢向顶极群落演替。群落内部的种间和种内竞争应是当前群落动态的主导因素。
黄云霞, 徐萱, 张莉芗, 宋玥, 骆争荣 (2016) 百山祖常绿阔叶林灌草层物种组成和 分布的10年动态. 生物多样性, 24, 1353-1363. DOI: 10.17520/biods.2016095.
Yunxia Huang, Xuan Xu, Lixiang Zhang, Yue Song, Zhengrong Luo (2016) Ten-years period of grass and small woody plant dynamics in a 5-ha evergreen forest plot in Baishanzu, Zhejiang Province. Biodiversity Science, 24, 1353-1363. DOI: 10.17520/biods.2016095.
生活型 Life form | 个体数 No. of individuals | 物种数 No. of species | Camargo指数 Camargo index | |||||
---|---|---|---|---|---|---|---|---|
2003 | 2013 | 2003 | 2013 | 2003 | 2013 | |||
乔木 Tree species | 9,402 | 8,052 | 29 | 26 | 0.224 | 0.173 | ||
直立灌木 Erective shrubs | 43,352 | 29,003 | 79 | 72 | 0.167 | 0.135 | ||
蔓性灌木 Spreading shrubs | 3,431 | 2,453 | 10 | 9 | 0.176 | 0.229 | ||
木质藤本 Woody lianas | 1,630 | 1,170 | 17 | 16 | 0.191 | 0.130 | ||
草质藤本 Herbaceous lianas | 26 | 6 | 2 | 2 | 0.692 | 1.000 | ||
草本蕨类 Herbaceous ferns | 21,333 | 16,323 | 6 | 6 | 0.198 | 0.225 | ||
其他草本 Other herbs | 6,549 | 4,090 | 37 | 35 | 0.145 | 0.192 | ||
总计 Total | 85,723 | 61,097 | 180 | 166 | 0.127 | 0.114 |
表1 2003-2013年百山祖常绿阔叶林灌草层各生活型的数量变化特征
Table 1 Shifts in numerical characteristics of different life forms in the shrub and grass layer of the Baishanzu evergreen broad- leaved forest between 2003 and 2013
生活型 Life form | 个体数 No. of individuals | 物种数 No. of species | Camargo指数 Camargo index | |||||
---|---|---|---|---|---|---|---|---|
2003 | 2013 | 2003 | 2013 | 2003 | 2013 | |||
乔木 Tree species | 9,402 | 8,052 | 29 | 26 | 0.224 | 0.173 | ||
直立灌木 Erective shrubs | 43,352 | 29,003 | 79 | 72 | 0.167 | 0.135 | ||
蔓性灌木 Spreading shrubs | 3,431 | 2,453 | 10 | 9 | 0.176 | 0.229 | ||
木质藤本 Woody lianas | 1,630 | 1,170 | 17 | 16 | 0.191 | 0.130 | ||
草质藤本 Herbaceous lianas | 26 | 6 | 2 | 2 | 0.692 | 1.000 | ||
草本蕨类 Herbaceous ferns | 21,333 | 16,323 | 6 | 6 | 0.198 | 0.225 | ||
其他草本 Other herbs | 6,549 | 4,090 | 37 | 35 | 0.145 | 0.192 | ||
总计 Total | 85,723 | 61,097 | 180 | 166 | 0.127 | 0.114 |
物种 Species | 株数 Number of individuals | 频度 Frequency | 重要值排序 Rank of IV | |||||
---|---|---|---|---|---|---|---|---|
2003 | 2013 | 2003 | 2013 | 2003 | 2013 | |||
木质藤本植物(前3位) Woody lianas (Top 3) | ||||||||
显脉野木瓜 Stauntonia conspicua | 994 (60.98%) | 914 (78.12%) | 474 (59.62%) | 524 (77.86%) | 1 | 1 | ||
钻地风 Schizophragma integrifolium | 264 (16.19%) | 148 (12.65%) | 119 (14.97%) | 68 (10.10%) | 2 | 2 | ||
南五味子 Kadsura longipedunculata | 167 (10.25%) | 21 (1.79%) | 83 (10.44%) | 16 (2.38%) | 3 | 3 | ||
乔木(前3位) Tree species (Top 3) | ||||||||
短尾柯 Lithocarpus brevicaudatus | 3,430 (36.48%) | 4,366 (54.22%) | 1,288 (26.62%) | 1,536 (38.79%) | 1 | 1 | ||
多脉青冈 Cyclobalanopsis multinervis | 1,716 (18.25%) | 1,200 (14.90%) | 885 (18.28%) | 747 (18.86%) | 2 | 2 | ||
木荷 Schima superba | 980 (10.42%) | 621 (7.71%) | 468 (9.67%) | 362 (9.18%) | 3 | 3 | ||
直立灌木(前5位) Erective shrubs (Top 5) | ||||||||
连蕊茶 Camellia cuspidata | 5,894 (13.59%) | 4,384 (19.38%) | 1,561 (13.18%) | 1,558 (16.07%) | 1 | 1 | ||
新木姜子 Neolitsea aurata | 5,856 (13.51%) | 4,318 (14.89%) | 1,517 (12.81%) | 1,501 (15.22%) | 2 | 2 | ||
薄叶山矾 Symplocos anomala | 4,560 (10.52%) | 1,691 (0.62%) | 973 (5.22%) | 875 (1.43%) | 3 | 14 | ||
叶萼山矾 Symplocos phyllocalyx | 795 (1.83%) | 4,426 (15.26%) | 142 (1.20%) | 1,117 (11.33%) | 15 | 3 | ||
四川山矾 Symplocos setchuensis | 4,032 (9.30%) | 2,912 (10.04%) | 990 (5.36%) | 1,000 (10.14%) | 4 | 4 | ||
厚叶红淡比 Cleyera pachyphylla | 4,824 (11.13%) | 2,114 (7.29%) | 525 (4.43%) | 592 (6.00%) | 5 | 5 | ||
蔓性/攀援灌木(前3位) Spreading shrubs (Top 3) | ||||||||
尖叶菝葜 Smilax arisanensis | 2,752 (80.21%) | 1,667 (67.96%) | 1,129 (81.46%) | 852 (72.63%) | 1 | 1 | ||
福建悬钩子 Rubus fujianensis | 434 (12.65%) | 579 (23.60%) | 133 (9.60%) | 159 (13.55%) | 2 | 2 | ||
巴东胡颓子 Elaeagnus difficilis | 95 (2.77%) | 36 (1.47%) | 50 (3.61%) | 32 (2.73%) | 3 | 5 | ||
暗色菝葜 Smilax lanceifolia var. opaca | 34 (0.99%) | 66 (2.69%) | 20 (1.44%) | 47 (4.01%) | 5 | 3 | ||
草本蕨类植物(前3位) Herbaceous ferns (Top 3) | ||||||||
华东瘤足蕨 Plagiogyria japonica | 20,106 (94.25%) | 14,477 (88.69%) | 1,683 (84.02%) | 1,621 (79.27%) | 1 | 1 | ||
美观复叶耳蕨 Arachniodes speciosa | 746 (3.50%) | 479 (2.93%) | 162 (8.09%) | 80 (3.91%) | 2 | 3 | ||
林下凸轴蕨 Metathelypteris hattorii | 286 (1.34%) | 1,143 (7.00%) | 96 (4.79%) | 289 (14.13%) | 3 | 2 | ||
草质藤本 Herbaceous lianas | ||||||||
鸡屎藤 Paederia foetida | 21 (80.77%) | 3 (50.00%) | 11 (68.75%) | 2 (66.67%) | 1 | 1 | ||
三叶崖爬藤 Tetrastigma hemsleyanum | 5 (19.23%) | 3 (50.00%) | 5 (31.25%) | 1 (33.33%) | 2 | 2 | ||
其他草本植物(前5位) Other herbs (Top 5) | ||||||||
薹草属多种 Carex spp. | 2,415 (36.88%) | 1,570 (38.39%) | 715 (45.60%) | 412 (40.95%) | 1 | 1 | ||
麦冬 Ophiopogon japonicus | 2,683 (40.97%) | 763 (18.65%) | 508 (32.40%) | 152 (15.11%) | 2 | 2 | ||
山麦冬 Liriope spicata | 185 (2.82%) | 27 (0.66%) | 46 (2.93%) | 15 (1.49%) | 3 | 11 | ||
求米草 Oplismenus undulatifolius | 36 (0.55%) | 637 (15.57%) | 5 (0.32%) | 22 (2.18%) | 18 | 3 | ||
异药花 Fordiophyton faberi | 111 (1.69%) | 333 (8.14%) | 36 (2.30%) | 50 (4.97%) | 4 | 4 | ||
赤车 Pellionia radicans | 139 (2.12%) | 240 (5.87%) | 23 (1.47%) | 42 (4.17%) | 5 | 5 |
表2 2003-2013年百山祖常绿阔叶林灌草层主要物种重要值(IV)的变化
Table 2 Shifts in importance value (IV) of dominant species in the shrub and grass layer of the Baishanzu evergreen broad-leaved forest between 2003 and 2013
物种 Species | 株数 Number of individuals | 频度 Frequency | 重要值排序 Rank of IV | |||||
---|---|---|---|---|---|---|---|---|
2003 | 2013 | 2003 | 2013 | 2003 | 2013 | |||
木质藤本植物(前3位) Woody lianas (Top 3) | ||||||||
显脉野木瓜 Stauntonia conspicua | 994 (60.98%) | 914 (78.12%) | 474 (59.62%) | 524 (77.86%) | 1 | 1 | ||
钻地风 Schizophragma integrifolium | 264 (16.19%) | 148 (12.65%) | 119 (14.97%) | 68 (10.10%) | 2 | 2 | ||
南五味子 Kadsura longipedunculata | 167 (10.25%) | 21 (1.79%) | 83 (10.44%) | 16 (2.38%) | 3 | 3 | ||
乔木(前3位) Tree species (Top 3) | ||||||||
短尾柯 Lithocarpus brevicaudatus | 3,430 (36.48%) | 4,366 (54.22%) | 1,288 (26.62%) | 1,536 (38.79%) | 1 | 1 | ||
多脉青冈 Cyclobalanopsis multinervis | 1,716 (18.25%) | 1,200 (14.90%) | 885 (18.28%) | 747 (18.86%) | 2 | 2 | ||
木荷 Schima superba | 980 (10.42%) | 621 (7.71%) | 468 (9.67%) | 362 (9.18%) | 3 | 3 | ||
直立灌木(前5位) Erective shrubs (Top 5) | ||||||||
连蕊茶 Camellia cuspidata | 5,894 (13.59%) | 4,384 (19.38%) | 1,561 (13.18%) | 1,558 (16.07%) | 1 | 1 | ||
新木姜子 Neolitsea aurata | 5,856 (13.51%) | 4,318 (14.89%) | 1,517 (12.81%) | 1,501 (15.22%) | 2 | 2 | ||
薄叶山矾 Symplocos anomala | 4,560 (10.52%) | 1,691 (0.62%) | 973 (5.22%) | 875 (1.43%) | 3 | 14 | ||
叶萼山矾 Symplocos phyllocalyx | 795 (1.83%) | 4,426 (15.26%) | 142 (1.20%) | 1,117 (11.33%) | 15 | 3 | ||
四川山矾 Symplocos setchuensis | 4,032 (9.30%) | 2,912 (10.04%) | 990 (5.36%) | 1,000 (10.14%) | 4 | 4 | ||
厚叶红淡比 Cleyera pachyphylla | 4,824 (11.13%) | 2,114 (7.29%) | 525 (4.43%) | 592 (6.00%) | 5 | 5 | ||
蔓性/攀援灌木(前3位) Spreading shrubs (Top 3) | ||||||||
尖叶菝葜 Smilax arisanensis | 2,752 (80.21%) | 1,667 (67.96%) | 1,129 (81.46%) | 852 (72.63%) | 1 | 1 | ||
福建悬钩子 Rubus fujianensis | 434 (12.65%) | 579 (23.60%) | 133 (9.60%) | 159 (13.55%) | 2 | 2 | ||
巴东胡颓子 Elaeagnus difficilis | 95 (2.77%) | 36 (1.47%) | 50 (3.61%) | 32 (2.73%) | 3 | 5 | ||
暗色菝葜 Smilax lanceifolia var. opaca | 34 (0.99%) | 66 (2.69%) | 20 (1.44%) | 47 (4.01%) | 5 | 3 | ||
草本蕨类植物(前3位) Herbaceous ferns (Top 3) | ||||||||
华东瘤足蕨 Plagiogyria japonica | 20,106 (94.25%) | 14,477 (88.69%) | 1,683 (84.02%) | 1,621 (79.27%) | 1 | 1 | ||
美观复叶耳蕨 Arachniodes speciosa | 746 (3.50%) | 479 (2.93%) | 162 (8.09%) | 80 (3.91%) | 2 | 3 | ||
林下凸轴蕨 Metathelypteris hattorii | 286 (1.34%) | 1,143 (7.00%) | 96 (4.79%) | 289 (14.13%) | 3 | 2 | ||
草质藤本 Herbaceous lianas | ||||||||
鸡屎藤 Paederia foetida | 21 (80.77%) | 3 (50.00%) | 11 (68.75%) | 2 (66.67%) | 1 | 1 | ||
三叶崖爬藤 Tetrastigma hemsleyanum | 5 (19.23%) | 3 (50.00%) | 5 (31.25%) | 1 (33.33%) | 2 | 2 | ||
其他草本植物(前5位) Other herbs (Top 5) | ||||||||
薹草属多种 Carex spp. | 2,415 (36.88%) | 1,570 (38.39%) | 715 (45.60%) | 412 (40.95%) | 1 | 1 | ||
麦冬 Ophiopogon japonicus | 2,683 (40.97%) | 763 (18.65%) | 508 (32.40%) | 152 (15.11%) | 2 | 2 | ||
山麦冬 Liriope spicata | 185 (2.82%) | 27 (0.66%) | 46 (2.93%) | 15 (1.49%) | 3 | 11 | ||
求米草 Oplismenus undulatifolius | 36 (0.55%) | 637 (15.57%) | 5 (0.32%) | 22 (2.18%) | 18 | 3 | ||
异药花 Fordiophyton faberi | 111 (1.69%) | 333 (8.14%) | 36 (2.30%) | 50 (4.97%) | 4 | 4 | ||
赤车 Pellionia radicans | 139 (2.12%) | 240 (5.87%) | 23 (1.47%) | 42 (4.17%) | 5 | 5 |
图1 不同生活型物种2003年的重要值与其10年间变化的关系。黑色三角形表示异常值,其旁边标注了对应物种的学名, 实线表示不含异常值的回归直线。
Fig. 1 Relationships between importance values of species with different life forms in 2003 and its changes from 2003 to 2013. Solid triangles indicate oddities labeled with the corresponding species names nearby. Solid lines display linear regression curves for data without or excluding oddities.
生境 Habitat | 与生境负相关的物种数 Number of species positively associated with habitats | 与生境正相关的物种数 Number of species negatively associated with habitats | |||||
---|---|---|---|---|---|---|---|
2003 | 2013 | 差异 Difference | 2003 | 2013 | 差异 Difference | ||
山谷 Gull | 7 | 5 | -2 | 5 | 11 | 6 | |
陡坡 Steep slope | 1 | 1 | 0 | 1 | 1 | 0 | |
缓坡 Mild slope | 1 | 0 | -1 | 7 | 4 | -3 | |
山脊 Ridge | 4 | 2 | -2 | 4 | 4 | 0 |
表3 百山祖常绿阔叶林灌草层2003和2013年间与生境相关的物种在各类生境中的分布
Table 3 Number of species with remarkable association to each habitat type of the Baishanzu evergreen broad-leaved forest in 2003 and 2013
生境 Habitat | 与生境负相关的物种数 Number of species positively associated with habitats | 与生境正相关的物种数 Number of species negatively associated with habitats | |||||
---|---|---|---|---|---|---|---|
2003 | 2013 | 差异 Difference | 2003 | 2013 | 差异 Difference | ||
山谷 Gull | 7 | 5 | -2 | 5 | 11 | 6 | |
陡坡 Steep slope | 1 | 1 | 0 | 1 | 1 | 0 | |
缓坡 Mild slope | 1 | 0 | -1 | 7 | 4 | -3 | |
山脊 Ridge | 4 | 2 | -2 | 4 | 4 | 0 |
图2 灌草层内主要物种与生境因子间的典范对应分析排序图。黑点表示2003年的物种, 三角形表示2013年的物种。图中elev, slope, convex和BA分别表示10 m × 10 m样方的相对高度、坡度、凹凸度和其中胸径1 cm以上所有植物的胸高断面积。
Fig. 2 Ordination of the main species and four habitat factors in the canonical correspondence analysis (CCA). Black points indicate species living in 2003, while hollow triangles indicate species living in 2013. The marks of elev, slope and convex mean elevation, slope and convexity of 10 m × 10 m quadrates which plants was allocated in, while BA means basal area of all plants with DBH ≥ 1 cm in the quadrates.
生境 Habitat | 胸高断面积 Basal area (m2/quadrat) | 密度 Density (ind./m2) | Shannon-Wiener指数 Shannon-Wiener index | Simpson指数 Simpson index | |||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|
2003 | 2013 | 2003 | 2013 | 差异 Difference | 2003 | 2013 | 差异 Difference | ||||||
山谷 Gull | 1.36 | 1.54 | 1.08 | 2.44 | 2.29 | -0.15* | 0.86 | 0.84 | -0.02* | ||||
陡坡 Steep slope | 1.74 | 2.04 | 1.74 | 2.49 | 2.41 | -0.08 | 0.85 | 0.84 | -0.01 | ||||
缓坡 Mild slope | 1.82 | 1.80 | 1.25 | 2.41 | 2.25 | -0.16* | 0.85 | 0.84 | -0.01 | ||||
山脊 Ridge | 1.97 | 1.83 | 1.28 | 2.38 | 2.27 | -0.11* | 0.84 | 0.85 | 0.01 |
表4 百山祖常绿阔叶林各类生境的平均胸高断面积与2003-2013年灌草层植物密度和多样性的变化
Table 4 Average basal area and changes in density and diversity in each habitat in the Baishanzu evergreen broad-leaved forest between 2003 and 2013
生境 Habitat | 胸高断面积 Basal area (m2/quadrat) | 密度 Density (ind./m2) | Shannon-Wiener指数 Shannon-Wiener index | Simpson指数 Simpson index | |||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|
2003 | 2013 | 2003 | 2013 | 差异 Difference | 2003 | 2013 | 差异 Difference | ||||||
山谷 Gull | 1.36 | 1.54 | 1.08 | 2.44 | 2.29 | -0.15* | 0.86 | 0.84 | -0.02* | ||||
陡坡 Steep slope | 1.74 | 2.04 | 1.74 | 2.49 | 2.41 | -0.08 | 0.85 | 0.84 | -0.01 | ||||
缓坡 Mild slope | 1.82 | 1.80 | 1.25 | 2.41 | 2.25 | -0.16* | 0.85 | 0.84 | -0.01 | ||||
山脊 Ridge | 1.97 | 1.83 | 1.28 | 2.38 | 2.27 | -0.11* | 0.84 | 0.85 | 0.01 |
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