生物多样性 ›› 2015, Vol. 23 ›› Issue (2): 149-156.doi: 10.17520/biods.2014137

• • 上一篇    下一篇

亚热带常绿阔叶林林窗物种丰富度的影响因素

刘何铭1, 2, 杨庆松1, 2, 方晓峰1, 2, 马遵平1, 2, 沈国春1, 2, 张志国1, 2, 王樟华3, 王希华1, 2, *()   

  1. 1 浙江天童森林生态系统国家野外科学观测研究站, 浙江宁波 315114
    2 华东师范大学生态与环境科学学院, 上海 200241
    3 中国科学院上海辰山植物科学研究中心, 上海 201602
  • 收稿日期:2014-06-30 接受日期:2014-09-28 出版日期:2015-03-20
  • 通讯作者: 王希华 E-mail:xhwang@des.ecnu.edu.cn
  • 基金项目:
    国家自然科学基金重大国际合作项目(31210103920)

Influences on gap species richness in a subtropical evergreen broad- leaved forest

Heming Liu1, 2, Qingsong Yang1, 2, Xiaofeng Fang1, 2, Zunping Ma1, 2, Guochun Shen1, 2, Zhiguo Zhang1, 2, Zhanghua Wang3, Xihua Wang1, 2, *()   

  1. 1 Tiantong National Forest Ecosystem Observation and Research Station, Ningbo, Zhejiang 315114
    2 School of Ecological and Environmental Sciences, East China Normal University, Shanghai 200241
    3 Shanghai Chenshan Plant Science Research Center, Chinese Academy of Sciences, Shanghai 201602
  • Received:2014-06-30 Accepted:2014-09-28 Online:2015-03-20
  • Contact: Wang Xihua E-mail:xhwang@des.ecnu.edu.cn

林窗是森林群落物种多样性维持中十分重要的结构, 但其本身物种多样性的维持机制尚不清楚, 可能与相邻群落的物种丰富度、林窗面积以及土壤和地形因子等有关。本文选取浙江天童20 ha森林动态监测样地内的84个冠林窗作为研究对象, 分析了林窗内部以及不同距离邻域内的物种组成, 选取包含大部分林窗内物种(≥75%), 并且对林窗内物种丰富度影响最大的邻域作为相关邻域(林窗周围成熟植株通过种子传播影响林窗物种丰富度, 把距离冠林窗边缘5 m范围内的区域作为种源效应邻域; 由于距离林窗较近, 物种组成与林窗形成时仍存活个体的物种组成相似, 故把距离冠林窗边缘4 m范围内的区域作为距离效应邻域), 然后计算种源效应邻域内成熟植株的物种丰富度、距离效应邻域内所有植株的物种丰富度、冠林窗的面积、地形因子(海拔、坡度、坡向、凸度)和土壤因子(pH值、全氮、全碳、全磷), 最后对11个潜在的影响因素进行随机组合, 构建广义线性回归模型, 筛选最优模型(AICc最小)进行分析。结果显示: 种源效应邻域内成熟植株的物种丰富度和冠林窗面积显著影响林窗内物种丰富度, 其他因素并未对冠林窗内物种丰富度产生显著影响。上述结果说明, 冠林窗面积的大小以及外部种源的丰富程度是决定冠林窗内物种丰富度的主要因素, 其他因素可能只产生间接影响。

关键词: 林窗大小, 物种多样性, 种源, 冠林窗面积, 天童, 森林群落, 动态样地

Canopy gaps, created by the death or disturbance of a mature tree, influence forest community structure. The opening of the canopy creates opportunity for species recruitment influencing species diversity in forest communities. However, the mechanisms that lead to species recruitment and the resulting diversity are still unclear. Factors that may affect species richness of canopy gaps include the gap area, species richness of adjacent areas (SSAA and SCAA represent the adjacent area where is the seed source of canopy gap and the adjacent area where species composition is spatially correlated with survivors in the canopy gap, respectively), topography, and soil nutrients. To test which factors significantly affect species richness, 11 parameters (canopy gap area, species richness of mature individuals in SSAA, species richness of all individuals in SCAA, slope, aspect, convexity, mean elevation, pH value, total nitrogen, total carbon, total phosphorus of each gap) were calculated for 84 canopy gaps in 20 ha Tiantong Dynamics Plot. The effect of SSAA and SCAA were quantified by species richness of mature individuals and richness of all free standing trees (DBH ≥ 1 cm) within 5 m and 4 m away from the edge of the gap, respectively. The size of adjacent area was defined as the area containing most of species present in the gap (≥75%) and having most significant effect on gap species richness. A generalized linear regression model was constructed for each random combinations of any subsets of 11 gap parameters. Our best model with lowest AICc showed that: only species richness of mature individuals in SSAA and the canopy gap area had a significant effect on the gap species richness. This result suggests that gap size and species richness of the seed source can directly influence species richness of canopy gaps. Additional factors may have an indirect influence and should be explored further.

Key words: gap size, species diversity, seed source, canopy gap area, Tiantong, forest community, dynamics plot

图1

冠林窗及扩展林窗分布。图中阴影部分代表冠林窗, 黑色实线为扩展林窗, 虚线为调查样线, 灰色实线为等高线。"

图2

不同尺度邻域内物种组成对冠林窗物种丰富度的影响程度(AIC)以及包含冠林窗内物种的比例。距离指冠林窗边缘向外扩展的距离。(a)所有植株; (b)成熟植株。"

图3

邻域物种丰富度对冠林窗内物种丰富度的影响(广义线性回归模型)。(a)距离效应邻域内所有植株的物种丰富度; (b)种源效应邻域内成熟植株的物种丰富度。实线为拟合曲线, 虚线内为预测值95%的置信区间。"

表1

影响冠林窗物种丰富度的主要因素"

参数
Parameter
OR值
Odds ratio
95%置信区间
95% of confidence interval
权重
Akaike weight (AICc)
种源效应邻域内成熟植株的物种丰富度
Species richness of mature individuals in SSAA
1.033 1.016-1.050 0.866
冠林窗的面积 Canopy gap size (log) 1.251 1.068-1.465 0.864
海拔 Elevation 1.001 0.999-1.004 0.428
距离效应邻域内所有植株的物种丰富度
Species richness of all individuals in SCAA
/ / 0.447
坡向 Aspect (cos(π-x) + 1.1) / / 0.377
pH值 pH / / 0.297
坡度 Slope / / 0.295
全氮 Total nitrogen / / 0.264
凸度 Convexity / / 0.264
全碳 Total carbon / / 0.256
全磷 Total phosphorus / / 0.253

附表1

浙江天童20 ha森林动态监测样地中冠林窗以及相邻群落物种组成名录"

种名
Species
先锋种或非先锋种
Pioneer species or non-pioneer species
种名
Species
先锋种或非先锋种
Pioneer species or non-pioneer species
锐角槭 Acer acutum - 厚壳树 Ehretia acuminata +
三角槭 Acer buergerianum - 中华杜英 Elaeocarpus chinensis -
毛脉槭 Acer pubinerve - 杜英 Elaeocarpus decipiens -
毛八角枫 Alangium kurzii + 中华卫矛 Euonymus nitidus -
赤杨叶 Alniphyllum fortunei + 细枝柃 Eurya loquaiana -
糙叶树 Aphananthe aspera + 格药柃 Eurya muricata -
棘茎楤木 Aralia echinocaulis + 窄基红褐柃 Eurya rubiginosa var. attenuata -
老鸦糊 Callicarpa giraldii - 野鸦椿 Euscaphis japonica -
秃红紫珠 Callicarpa rubella var. subglabra + 矮小天仙果 Ficus erecta -
毛花连蕊茶 Camellia fraterna - 苦枥木 Fraxinus insularis -
山茶 Camellia japonica - 银杏 Ginkgo biloba -
Camellia sinensis - 山桐子 Idesia polycarpa +
雷公鹅耳枥 Carpinus viminea - 短梗冬青 Ilex buergeri -
米槠 Castanopsis carlesii - 冬青 Ilex chinensis -
栲树 Castanopsis fargesii - 细刺枸骨 Ilex hylonoma -
苦槠 Castanopsis sclerophylla - 皱柄冬青 Ilex kengii -
紫弹树 Celtis biondii + 大叶冬青 Ilex latifolia -
朴树 Celtis sinensis + 小果冬青 Ilex micrococca -
西川朴 Celtis vandervoetiana + 铁冬青 Ilex rotunda -
迎春樱桃 Cerasus discoidea - 红毒茴 Illicium lanceolatum -
大叶早樱 Cerasus subhirtella - 胡桃楸 Juglans mandshurica -
南酸枣 Choerospondias axillaris - 刺楸 Kalopanax septemlobus +
香桂 Cinnamomum subavenium - 腺叶桂樱 Laurocerasus phaeosticta -
大青 Clerodendrum cyrtophyllum + 刺叶桂樱 Laurocerasus spinulosa -
红淡比 Cleyera japonica - 小蜡 Ligustrum sinense -
杉木 Cunninghamia lanceolata - 红果山胡椒 Lindera erythrocarpa +
赤皮青冈 Cyclobalanopsis gilva - 山胡椒 Lindera glauca +
青冈 Cyclobalanopsis glauca - 红脉钓樟 Lindera rubronervia +
细叶青冈 Cyclobalanopsis gracilis - 枫香树 Liquidambar formosana -
小叶青冈 Cyclobalanopsis myrsinaefolia - 港柯 Lithocarpus harlandii -
云山青冈 Cyclobalanopsis sessilifolia - 豹皮樟 Litsea coreana var. sinensis -
青钱柳 Cyclocarya paliurus + 山鸡椒 Litsea cubeba +
黄檀 Dalbergia hupeana - 黄丹木姜子 Litsea elongata -
虎皮楠 Daphniphyllum oldhami - 檵木 Loropetalum chinense +
山柿 Diospyros japonica - 薄叶润楠 Machilus leptophylla -
油柿 Diospyros oleifera - 红楠 Machilus thunbergii -
杨梅叶蚊母树 Distylium myricoides - 构棘 Maclura cochinchinensis -
天目木兰 Magnolia amoena - 鸡仔木 Sinoadina racemosa -
白背叶 Mallotus apelta + 赛山梅 Styrax confusus -
野桐 Mallotus japonicus var. floccosus + 薄叶山矾 Symplocos anomala -
红枝柴 Meliosma oldhamii - 总状山矾 Symplocos botryantha -
黐花 Mussaenda esquirolii + 黄牛奶树 Symplocos cochinchinensis var. laurina -
杨梅 Myrica rubra - 光叶山矾 Symplocos lancifolia -
浙江新木姜子 Neolitsea aurata var. chekiangensis - 光亮山矾 Symplocos lucida -
宁波木犀 Osmanthus cooperi - 老鼠矢 Symplocos stellaris -
木犀 Osmanthus fragrans - 山矾 Symplocos sumuntia -
紫楠 Phoebe sheareri - 赤楠 Syzygium buxifolium -
光叶石楠 Photinia glabra - 厚皮香 Ternstroemia gymnanthera -
青灰叶下珠 Phyllanthus glaucus + 榧树 Torreya grandis -
海金子 Pittosporum illicioides - 野漆树 Toxicodendron succedaneum +
化香树 Platycarya strobilacea + 木蜡树 Toxicodendron sylvestre -
豆腐柴 Premna microphylla - 棕榈 Trachycarpus fortunei -
石斑木 Rhaphiolepis indica - 山油麻 Trema cannabina var. dielsiana +
马银花 Rhododendron ovatum - 杭州榆 Ulmus changii -
杜鹃 Rhododendron simsii + 南烛 Vaccinium bracteatum -
盐肤木 Rhus chinensis + 江南越桔 Vaccinium mandarinorum -
无患子 Sapindus saponaria - 刺毛越桔 Vaccinium trichocladum -
檫木 Sassafras tzumu + 油桐 Vernicia fordii +
木荷 Schima superba - 宜昌荚蒾 Viburnum erosum -
青皮木 Schoepfia jasminodora + 大叶榉树 Zelkova schneideriana +
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