植物生态学报 ›› 2015, Vol. 39 ›› Issue (10): 990-1002.DOI: 10.17521/cjpe.2015.0096
黄永涛1,2, 姚兰1, 艾训儒1, 吕世安3, 丁易2,4,*()
出版日期:
2015-10-01
发布日期:
2015-10-24
通讯作者:
丁易
作者简介:
# 共同第一作者
基金资助:
HUANG Yong-Tao1,2, YAO Lan1, AI Xun-Ru1, LÜ Shi-An3, DING Yi2,4,*()
Online:
2015-10-01
Published:
2015-10-24
Contact:
Yi DING
About author:
# Co-first authors
摘要:
为深入研究亚热带常绿落叶阔叶混交林的生物多样性维持机制, 合理保护与利用此类森林植被, 以鄂西南两个自然保护区——星斗山和木林子典型的亚热带常绿落叶阔叶混交林为研究对象, 在野外样方调查的基础上进行了群落数量分类和排序, 分析了常绿和落叶物种比例随群落类型及环境因子变化的规律。研究结果表明: (1)将鄂西南两个自然保护区的亚热带常绿落叶阔叶混交林划分为5个群落类型: 即青冈-翅柃+尖连蕊茶(Cyclobalanopsis glauca-Eurya alata + Camellia cuspidata)群落(I)、川陕鹅耳枥+青冈-翅柃(Carpinus fargesiana + Cyclobalanopsis glauca-Eurya alata)群落(II)、川陕鹅耳枥+青冈-水马桑(Carpinus fargesiana + Cyclobalanopsis glauca-Weigela japonica)群落(III)、光叶水青冈+多脉青冈-翅柃(Fagus lucida + Cyclobalanopsis multinervis-Eurya alata)群落(IV)和川陕鹅耳枥+交让木+包果柯-翅柃(Carpinus fargesiana + Daphniphyllum macropodum + Lithocarpus cleistocarpus-Eurya alata)群落(V); (2)常绿和落叶物种的丰富度、多度、胸高断面积及重要值等指标随群落类型而变化。在群落II、III和V中落叶物种占优势, 而在群落I和IV中二者无显著差异; (3)各类型群落在小径级(1-5 cm)内落叶物种的丰富度大于常绿物种, 但多度、胸高断面积和重要值大都小于常绿物种。在中径级(5-10 cm)内, 群落I和群落IV的各项指标都表现为常绿物种大于落叶物种, 而群落II、群落III和群落V的则相反。在大径级(≥ 10 cm)内, 各群落类型的落叶物种都占显著优势, 其各项指标大于常绿物种; (4)海拔、坡度、坡向、土壤氮含量和土壤磷含量是影响各群落类型内常绿和落叶物种比例变化的主导因子。
黄永涛, 姚兰, 艾训儒, 吕世安, 丁易. 鄂西南两个自然保护区亚热带常绿落叶阔叶混交林类型及其常绿和落叶物种组成结构分析. 植物生态学报, 2015, 39(10): 990-1002. DOI: 10.17521/cjpe.2015.0096
HUANG Yong-Tao,YAO Lan,AI Xun-Ru,LÜ Shi-An,DING Yi. Quantitative classification of the subtropical evergreen-deciduous broadleaved mixed forest and the deciduous and evergreen species composition structure across two national nature reserves in the southwest of Hubei, China. Chinese Journal of Plant Ecology, 2015, 39(10): 990-1002. DOI: 10.17521/cjpe.2015.0096
图1 鄂西南两个保护区亚热带常绿落叶阔叶混交林92个样地的TWINSPAN分类树状图。D, 分类次数; N, 样方数。
Fig. 1 Dendrogram of TWINSPAN classification for 92 plots in the subtropical evergreen-deciduous broadleaved mixed forest across two nature reserves in Southwest Hubei, China. D, division number; N, number of plot.
图2 鄂西南两个保护区亚热带常绿落叶阔叶混交林92个样地的DCA二维排序图。罗马数字(I, II, III, IV, V)代表不同的群落类型。
Fig. 2 Two-dimensional DCA ordination diagram for 92 plots in the subtropical evergreen-deciduous broadleaved mixed forest across two nature reserves in Southwest Hubei, China. Roman numerals (I, II, III, IV, V) indicate different community types.
群落类型 Community type | 物种 Species | 多度 Abundance | 胸径 DBH (cm) | 重要值 IV | 叶习性 Leaf habit |
---|---|---|---|---|---|
群落I | 青冈 Cyclobalanopsis glauca | 752 | 6.38 | 2.48 | 常绿 Evergreen |
Type I | 翅柃 Eurya alata | 599 | 4.02 | 1.12 | 常绿 Evergreen |
尖连蕊茶 Camellia cuspidata | 487 | 3.64 | 1.11 | 常绿 Evergreen | |
木荷 Schima superba | 248 | 5.94 | 0.99 | 常绿 Evergreen | |
山矾 Symplocos sumuntia | 535 | 3.05 | 0.95 | 常绿 Evergreen | |
锥栗 Castanea henryi | 17 | 53.96 | 0.83 | 落叶 Deciduous | |
黄丹木姜子 Litsea elongata | 385 | 2.85 | 0.82 | 常绿 Evergreen | |
薄叶山矾 Symplocos anomala | 207 | 3.59 | 0.39 | 常绿 Evergreen | |
茶条果 Symplocos phyllocalyx | 137 | 3.81 | 0.38 | 常绿 Evergreen | |
四照花 Dendrobenthamia japonica | 70 | 7.58 | 0.36 | 落叶 Deciduous | |
群落II | 川陕鹅耳枥 Carpinus fargesiana | 669 | 4.05 | 1.28 | 落叶 Deciduous |
Type II | 山矾 Symplocos sumuntia | 1 007 | 2.57 | 1.26 | 常绿 Evergreen |
青冈 Cyclobalanopsis glauca | 382 | 4.84 | 1.02 | 常绿 Evergreen | |
四照花 Dendrobenthamia japonica | 367 | 3.9 | 0.84 | 落叶 Deciduous | |
亮叶桦 Betula luminifera | 92 | 15.45 | 0.78 | 落叶 Deciduous | |
翅柃 Eurya alata | 452 | 2.44 | 0.65 | 常绿 Evergreen | |
水马桑 Weigela japonica | 172 | 4.51 | 0.40 | 落叶 Deciduous | |
灯台树 Bothrocaryum controversum | 68 | 8.54 | 0.36 | 落叶 Deciduous | |
黄丹木姜子 Litsea elongata | 136 | 3.13 | 0.34 | 常绿 Evergreen | |
枫香树 Liquidambar formosana | 56 | 10.92 | 0.31 | 落叶 Deciduous | |
群落III | 川陕鹅耳枥 Carpinus fargesiana | 1 108 | 4.03 | 1.63 | 落叶 Deciduous |
Type III | 水马桑 Weigela japonica | 843 | 3.89 | 1.41 | 落叶 Deciduous |
青冈 Cyclobalanopsis glauca | 850 | 3.93 | 1.18 | 常绿 Evergreen | |
四照花 Dendrobenthamia japonica | 580 | 3.25 | 0.82 | 落叶 Deciduous | |
灰柯 Lithocarpus henryi | 353 | 4.90 | 0.73 | 常绿 Evergreen | |
亮叶桦 Betula luminifera | 144 | 9.80 | 0.72 | 落叶 Deciduous | |
小叶青冈 Cyclobalanopsis myrsinifolia | 432 | 4.76 | 0.68 | 常绿 Evergreen | |
盐肤木 Rhus chinensis | 163 | 7.14 | 0.57 | 落叶 Deciduous | |
贵定桤叶树 Clethra cavaleriei | 355 | 3.11 | 0.57 | 落叶 Deciduous | |
山鸡椒 Litsea cubeba | 344 | 3.08 | 0.57 | 落叶 Deciduous | |
群落IV | 光叶水青冈 Fagus lucida | 325 | 9.77 | 1.64 | 常绿 Evergreen |
Type IV | 多脉青冈 Cyclobalanopsis multinervis | 414 | 6.87 | 1.23 | 落叶 Deciduous |
翅柃 Eurya alata | 526 | 3.13 | 0.95 | 常绿 Evergreen | |
川陕鹅耳枥 Carpinus fargesiana | 140 | 8.16 | 0.62 | 落叶 Deciduous | |
宜昌润楠 Machilus ichangensis | 77 | 7.88 | 0.31 | 常绿 Evergreen | |
交让木 Daphniphyllum macropodum | 72 | 7.12 | 0.28 | 常绿 Evergreen | |
杜鹃 Rhododendron simsii | 196 | 2.11 | 0.20 | 常绿 Evergreen | |
荚蒾 Viburnum dilatatum | 119 | 2.04 | 0.20 | 落叶 Deciduous | |
光叶山矾 Symplocos lancifolia | 60 | 4.28 | 0.17 | 常绿 Evergreen | |
包果柯 Lithocarpus cleistocarpus | 48 | 7.69 | 0.17 | 常绿 Evergreen | |
群落V | 川陕鹅耳枥 Carpinus fargesiana | 1 359 | 7.63 | 4.67 | 落叶 Deciduous |
Type V | 翅柃 Eurya alata | 1 710 | 2.84 | 2.53 | 常绿 Evergreen |
包果柯 Lithocarpus cleistocarpus | 502 | 8.13 | 1.71 | 常绿 Evergreen | |
交让木 Daphniphyllum macropodum | 461 | 5.08 | 1.53 | 常绿 Evergreen | |
亮叶桦 Betula luminifera | 201 | 12.09 | 1.40 | 落叶 Deciduous | |
齿缘吊钟花 Enkianthus serrulatus | 1 268 | 3.03 | 1.37 | 落叶 Deciduous | |
长蕊杜鹃 Rhododendron stamineum | 854 | 3.29 | 1.17 | 常绿 Evergreen | |
水马桑 Weigela japonica | 345 | 5.56 | 1.17 | 落叶 Deciduous | |
锥栗 Castanea henryi | 145 | 16.32 | 1.16 | 落叶 Deciduous | |
石灰花楸 Sorbus folgneri | 519 | 4.91 | 1.09 | 落叶 Deciduous |
表1 鄂西南两个保护区亚热带常绿落叶阔叶混交林5种群落类型内重要值排名前10位的物种
Table 1 Importance value ( IV ) of the top 10 species in the five community types in the subtropical evergreen-deciduous broadleaved mixed forest across two nature reserves in Southwest Hubei, China
群落类型 Community type | 物种 Species | 多度 Abundance | 胸径 DBH (cm) | 重要值 IV | 叶习性 Leaf habit |
---|---|---|---|---|---|
群落I | 青冈 Cyclobalanopsis glauca | 752 | 6.38 | 2.48 | 常绿 Evergreen |
Type I | 翅柃 Eurya alata | 599 | 4.02 | 1.12 | 常绿 Evergreen |
尖连蕊茶 Camellia cuspidata | 487 | 3.64 | 1.11 | 常绿 Evergreen | |
木荷 Schima superba | 248 | 5.94 | 0.99 | 常绿 Evergreen | |
山矾 Symplocos sumuntia | 535 | 3.05 | 0.95 | 常绿 Evergreen | |
锥栗 Castanea henryi | 17 | 53.96 | 0.83 | 落叶 Deciduous | |
黄丹木姜子 Litsea elongata | 385 | 2.85 | 0.82 | 常绿 Evergreen | |
薄叶山矾 Symplocos anomala | 207 | 3.59 | 0.39 | 常绿 Evergreen | |
茶条果 Symplocos phyllocalyx | 137 | 3.81 | 0.38 | 常绿 Evergreen | |
四照花 Dendrobenthamia japonica | 70 | 7.58 | 0.36 | 落叶 Deciduous | |
群落II | 川陕鹅耳枥 Carpinus fargesiana | 669 | 4.05 | 1.28 | 落叶 Deciduous |
Type II | 山矾 Symplocos sumuntia | 1 007 | 2.57 | 1.26 | 常绿 Evergreen |
青冈 Cyclobalanopsis glauca | 382 | 4.84 | 1.02 | 常绿 Evergreen | |
四照花 Dendrobenthamia japonica | 367 | 3.9 | 0.84 | 落叶 Deciduous | |
亮叶桦 Betula luminifera | 92 | 15.45 | 0.78 | 落叶 Deciduous | |
翅柃 Eurya alata | 452 | 2.44 | 0.65 | 常绿 Evergreen | |
水马桑 Weigela japonica | 172 | 4.51 | 0.40 | 落叶 Deciduous | |
灯台树 Bothrocaryum controversum | 68 | 8.54 | 0.36 | 落叶 Deciduous | |
黄丹木姜子 Litsea elongata | 136 | 3.13 | 0.34 | 常绿 Evergreen | |
枫香树 Liquidambar formosana | 56 | 10.92 | 0.31 | 落叶 Deciduous | |
群落III | 川陕鹅耳枥 Carpinus fargesiana | 1 108 | 4.03 | 1.63 | 落叶 Deciduous |
Type III | 水马桑 Weigela japonica | 843 | 3.89 | 1.41 | 落叶 Deciduous |
青冈 Cyclobalanopsis glauca | 850 | 3.93 | 1.18 | 常绿 Evergreen | |
四照花 Dendrobenthamia japonica | 580 | 3.25 | 0.82 | 落叶 Deciduous | |
灰柯 Lithocarpus henryi | 353 | 4.90 | 0.73 | 常绿 Evergreen | |
亮叶桦 Betula luminifera | 144 | 9.80 | 0.72 | 落叶 Deciduous | |
小叶青冈 Cyclobalanopsis myrsinifolia | 432 | 4.76 | 0.68 | 常绿 Evergreen | |
盐肤木 Rhus chinensis | 163 | 7.14 | 0.57 | 落叶 Deciduous | |
贵定桤叶树 Clethra cavaleriei | 355 | 3.11 | 0.57 | 落叶 Deciduous | |
山鸡椒 Litsea cubeba | 344 | 3.08 | 0.57 | 落叶 Deciduous | |
群落IV | 光叶水青冈 Fagus lucida | 325 | 9.77 | 1.64 | 常绿 Evergreen |
Type IV | 多脉青冈 Cyclobalanopsis multinervis | 414 | 6.87 | 1.23 | 落叶 Deciduous |
翅柃 Eurya alata | 526 | 3.13 | 0.95 | 常绿 Evergreen | |
川陕鹅耳枥 Carpinus fargesiana | 140 | 8.16 | 0.62 | 落叶 Deciduous | |
宜昌润楠 Machilus ichangensis | 77 | 7.88 | 0.31 | 常绿 Evergreen | |
交让木 Daphniphyllum macropodum | 72 | 7.12 | 0.28 | 常绿 Evergreen | |
杜鹃 Rhododendron simsii | 196 | 2.11 | 0.20 | 常绿 Evergreen | |
荚蒾 Viburnum dilatatum | 119 | 2.04 | 0.20 | 落叶 Deciduous | |
光叶山矾 Symplocos lancifolia | 60 | 4.28 | 0.17 | 常绿 Evergreen | |
包果柯 Lithocarpus cleistocarpus | 48 | 7.69 | 0.17 | 常绿 Evergreen | |
群落V | 川陕鹅耳枥 Carpinus fargesiana | 1 359 | 7.63 | 4.67 | 落叶 Deciduous |
Type V | 翅柃 Eurya alata | 1 710 | 2.84 | 2.53 | 常绿 Evergreen |
包果柯 Lithocarpus cleistocarpus | 502 | 8.13 | 1.71 | 常绿 Evergreen | |
交让木 Daphniphyllum macropodum | 461 | 5.08 | 1.53 | 常绿 Evergreen | |
亮叶桦 Betula luminifera | 201 | 12.09 | 1.40 | 落叶 Deciduous | |
齿缘吊钟花 Enkianthus serrulatus | 1 268 | 3.03 | 1.37 | 落叶 Deciduous | |
长蕊杜鹃 Rhododendron stamineum | 854 | 3.29 | 1.17 | 常绿 Evergreen | |
水马桑 Weigela japonica | 345 | 5.56 | 1.17 | 落叶 Deciduous | |
锥栗 Castanea henryi | 145 | 16.32 | 1.16 | 落叶 Deciduous | |
石灰花楸 Sorbus folgneri | 519 | 4.91 | 1.09 | 落叶 Deciduous |
图3 鄂西南两个保护区亚热带常绿落叶阔叶混交林5种群落类型的常绿和落叶物种丰富度、多度、胸高断面积和重要值(平均值±标准偏差)。不同小写字母(a, b)表示显著差异(p < 0.05)。I、II、III、IV、V分别为青冈-翅柃+尖连蕊茶群落, 川陕鹅耳枥+青冈-翅柃群落, 川陕鹅耳枥+青冈-水马桑群落, 光叶水青冈+多脉青冈-翅柃群落和川陕鹅耳枥+交让木+包果柯-翅柃 群落。
Fig. 3 Species richness, abundance, basal area and importance value of the five forest types in the subtropical evergreen-deciduous broadleaved mixed forest across two nature reserves in Southwest Hubei, China (mean ± SD). Different lowercase letters (a, b) indicate significant differences at the level of 0.05. I, II, III, IV, V indicate different community types: Cyclobalanopsis glauca-Eurya alata + Camellia cuspidata, Carpinus fargesiana + Cyclobalanopsis glauca-Eurya alata, Carpinus fargesiana + Cyclobalanopsis glauca-Weigela japonica, Fagus lucida + Cyclobalanopsis multinervis-Eurya alata, Carpinus fargesiana + Daphniphyllum macropodum + Lithocarpus cleistocarpus-Eurya alata.
图4 鄂西南两个保护区亚热带常绿落叶阔叶混交林5种群落类型的常绿和落叶物种不同径级的物种丰富度、多度、胸高断面积和重要值比例。a, 胸径1-5 cm; b, 胸径5-10 cm; c, 胸径≥10 cm。I-V同图3。
Fig. 4 Evergreen/deciduous ratio of species richness, abundance, basal area and importance value with size class in subtropical evergreen-deciduous broadleaved mixed forest across two nature reserves in Southwest Hubei, China. a, DBH 1-5 cm; b, DBH 5-10 cm; c, DBH ≥10 cm; DBH, diameter at breast height. I-V see Fig. 3.
环境因子 Environmental variables | R2 | p |
---|---|---|
海拔 Elevation | 0.126 7 | 0.001*** |
坡度 Slope | 0.028 5 | 0.001*** |
坡向 Aspect | 0.002 4 | 0.014* |
土壤含水量 Soil water content | 0.001 7 | 0.071 |
pH | 0.002 9 | 0.059 |
土壤有机质 Soil organic matter | 0.000 7 | 0.092 |
土壤全氮 Total nitrogen | 0.001 1 | 0.002** |
土壤全磷 Total phosphorus | 0.036 4 | 0.001*** |
土壤碱解氮 Available nitrogen | 0.017 8 | 0.001*** |
土壤速效磷 Available phosphorus | 0.245 7 | 0.001*** |
表2 鄂西南两个保护区亚热带常绿落叶阔叶混交林内环境因子对物种分布的影响
Table 2 Effects of environmental factors on species distribution in subtropical evergreen-deciduous broadleaved mixed forest across two nature reserves in Southwest Hubei, China
环境因子 Environmental variables | R2 | p |
---|---|---|
海拔 Elevation | 0.126 7 | 0.001*** |
坡度 Slope | 0.028 5 | 0.001*** |
坡向 Aspect | 0.002 4 | 0.014* |
土壤含水量 Soil water content | 0.001 7 | 0.071 |
pH | 0.002 9 | 0.059 |
土壤有机质 Soil organic matter | 0.000 7 | 0.092 |
土壤全氮 Total nitrogen | 0.001 1 | 0.002** |
土壤全磷 Total phosphorus | 0.036 4 | 0.001*** |
土壤碱解氮 Available nitrogen | 0.017 8 | 0.001*** |
土壤速效磷 Available phosphorus | 0.245 7 | 0.001*** |
类型1) Type1) | 物种丰富度2) Species richness2) | 环境变量3) Environmental variables3) | 参数4) Parameter4) | |||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
E | S | A | SWC | pH | SOM | TN | TP | AN | AP | AIC | p | |||
I | ESR | -0.26 | -1.16 | 1.11 | -46.83 | 0.020* | ||||||||
DSR | -1.55 | -32.35 | 0.329 | |||||||||||
II | ESR | 1.26 | -4.31 | 2.54 | -0.95 | -55.14 | 0.008* | |||||||
DSR | -55.29 | 0.007* | ||||||||||||
III | ESR | -41.90 | 0.019* | |||||||||||
DSR | 7.85 | 0.29 | -0.68 | -64.30 | 0.032* | |||||||||
IV | ESR | -0.13 | 4.16 | 1.96 | -11.01 | 0.021* | ||||||||
DSR | -1.98 | -24.49 | 0.053 | |||||||||||
V | ESR | -0.67 | -0.57 | -107.40 | 0.022* | |||||||||
DSR | -2.32 | -129.68 | 0.005* |
表3 鄂西南两个保护区亚热带常绿落叶阔叶混交林5种群落常绿和落叶物种丰富度与环境因子的多元逐步回归分析
Table 3 Multiple regression analysis between species richness and environmental variables in the five forest types in subtropical evergreen-deciduous broadleaved mixed forest across two nature reserves in Southwest Hubei, China
类型1) Type1) | 物种丰富度2) Species richness2) | 环境变量3) Environmental variables3) | 参数4) Parameter4) | |||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
E | S | A | SWC | pH | SOM | TN | TP | AN | AP | AIC | p | |||
I | ESR | -0.26 | -1.16 | 1.11 | -46.83 | 0.020* | ||||||||
DSR | -1.55 | -32.35 | 0.329 | |||||||||||
II | ESR | 1.26 | -4.31 | 2.54 | -0.95 | -55.14 | 0.008* | |||||||
DSR | -55.29 | 0.007* | ||||||||||||
III | ESR | -41.90 | 0.019* | |||||||||||
DSR | 7.85 | 0.29 | -0.68 | -64.30 | 0.032* | |||||||||
IV | ESR | -0.13 | 4.16 | 1.96 | -11.01 | 0.021* | ||||||||
DSR | -1.98 | -24.49 | 0.053 | |||||||||||
V | ESR | -0.67 | -0.57 | -107.40 | 0.022* | |||||||||
DSR | -2.32 | -129.68 | 0.005* |
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