生物多样性 ›› 2016, Vol. 24 ›› Issue (6): 639-648. DOI: 10.17520/biods.2015134
刘晓静1, 任思远2, 李鹿鑫2, 叶永忠2, 袁志良2, 王婷2,,A;*()
收稿日期:
2016-02-15
接受日期:
2016-06-06
出版日期:
2016-06-20
发布日期:
2016-06-20
通讯作者:
王婷
基金资助:
Xiaojing Liu1, Siyuan Ren2, Luxin Li2, Yongzhong Ye2, Zhiliang Yuan2, Ting Wang2,*()
Received:
2016-02-15
Accepted:
2016-06-06
Online:
2016-06-20
Published:
2016-06-20
Contact:
Wang Ting
摘要:
密度制约对物种共存起着重要作用。随着密度制约效应研究的深入, 亲缘关系较近的物种间表现出的密度制约效应逐渐被人们认识。本研究基于2009和2014年对宝天曼1 ha落叶阔叶林样地的2次调查数据, 利用广义线性混合模型分析了重要值排名前11位的物种不同邻域尺度的密度制约效应对不同径级(小径级(1 cm ≤ DBH < 5 cm)、中径级(5 cm ≤ DBH < 10 cm)、大径级(DBH ≥ 10 cm))目标个体存活的影响。研究表明: (1) 5年间样地中DBH ≥ 1 cm的所有个体的年均死亡率和增员率分别为5.85%和0.27%; (2)有5个物种的个体存活率与同种邻体个体数及同种邻体胸高断面积显著负相关; (3)小径级个体的存活在5 m的邻域范围内受同种密度制约和谱系密度制约的影响都很显著; 中径级个体在3个尺度上受到的密度制约和谱系密度制约的影响都不大; 大径级个体在7.5 m、10 m邻域范围内受谱系密度制约影响显著。结果表明, 同种密度制约和谱系密度制约效应对宝天曼落叶阔叶林不同生长阶段的树木个体影响不同。
刘晓静, 任思远, 李鹿鑫, 叶永忠, 袁志良, 王婷 (2016) 宝天曼国家级自然保护区落叶阔叶林密度制约效应对树木存活的影响. 生物多样性, 24, 639-648. DOI: 10.17520/biods.2015134.
Xiaojing Liu, Siyuan Ren, Luxin Li, Yongzhong Ye, Zhiliang Yuan, Ting Wang (2016) Detecting density dependence on tree survival in a deciduous broad- leaved forest in Baotianman National Nature Reserve. Biodiversity Science, 24, 639-648. DOI: 10.17520/biods.2015134.
径级 DBH size | 物种数 Number of species | 个体数 Number of individuals | 死亡数(死亡率) Number of mortality (mortality rate, %) | 新增数(增员率) Number of recruitment (recruitment rate, %) | |||
---|---|---|---|---|---|---|---|
2009 | 2014 | 2009 | 2014 | ||||
小径级 Small DBH size (1 cm ≤ DBH < 5 cm) | 54 | 46 | 1,538 | 1,091 | 482 (31.34) | 35 (2.28) | |
中径级 Medium DBH size (5 cm ≤ DBH < 10 cm) | 39 | 35 | 600 | 523 | 197 (32.83) | 120 (20.0) | |
大径级 Large DBH size (DBH ≥ 10 cm) | 22 | 24 | 417 | 381 | 68 (16.31) | 32 (7.68) | |
总计 Total (DBH ≥ 1 cm) | 58 | 53 | 2,555 | 1,995 | 747 (29.24) | 187 |
表1 宝天曼国家级自然保护区1 ha落叶阔叶林样地2009和2014年两次调查概况
Table 1 Summary data of two census in a 1-ha deciduous broad-leaved forest plot in Baotianman National Nature Reserve in 2009 and 2014
径级 DBH size | 物种数 Number of species | 个体数 Number of individuals | 死亡数(死亡率) Number of mortality (mortality rate, %) | 新增数(增员率) Number of recruitment (recruitment rate, %) | |||
---|---|---|---|---|---|---|---|
2009 | 2014 | 2009 | 2014 | ||||
小径级 Small DBH size (1 cm ≤ DBH < 5 cm) | 54 | 46 | 1,538 | 1,091 | 482 (31.34) | 35 (2.28) | |
中径级 Medium DBH size (5 cm ≤ DBH < 10 cm) | 39 | 35 | 600 | 523 | 197 (32.83) | 120 (20.0) | |
大径级 Large DBH size (DBH ≥ 10 cm) | 22 | 24 | 417 | 381 | 68 (16.31) | 32 (7.68) | |
总计 Total (DBH ≥ 1 cm) | 58 | 53 | 2,555 | 1,995 | 747 (29.24) | 187 |
图1 宝天曼国家级自然保护区1 ha落叶阔叶林样地木本植物物种的谱系关系
Fig. 1 Phylogenetic relationship among woody species in a 1-ha deciduous broad-leaved forest plot in Baotianman National Nature Reserve
物种 Species | 拉丁名 Latin name | 重要值 IV | |
---|---|---|---|
2009 | 2014 | ||
槲栎 | Quercus aliena | 20.46 | 21.65 |
水榆花楸 | Sorbus alnifolia | 16.69 | 19.17 |
三桠乌药 | Lindera obtusiloba | 11.26 | 7.59 |
青榨槭 | Acer davidii | 9.62 | 10.88 |
野茉莉 | Styrax japonicus | 4.94 | 5.72 |
毛樱桃 | Cerasus tomentosa | 3.49 | 3.53 |
连翘 | Forsythia suspensa | 2.49 | 2.75 |
白檀 | Symplocos paniculata | 2.42 | 1.88 |
卫矛 | Euonymus alatus | 2.42 | 2.41 |
秦岭木姜子 | Litsea tsinlingensis | 2.38 | |
元宝槭 | Acer truncatum | 2.24 | 2.70 |
千金榆 | Carpinus cordata | 1.75 | 2.00 |
山梅花 | Philadelphus incanus | 1.59 | 1.35 |
暖木 | Meliosma veitchioru | 1.52 | 1.74 |
泡花树 | Meliosma cuneifolia | 1.44 | 1.59 |
马鞍树 | Maackia hupehensi | 1.55 | |
桦叶荚蒾 | Viburnum betulifolium | 1.32 | 1.59 |
四照花 | Dendrobenthamia japonica var. chinensis | 1.25 | 1.55 |
华东椴 | Tilia japonica | 1.06 | |
老鸹铃 | Styrax hemsleyanus | 1.02 | |
灯台树 | Bothrocaryum controversum | 0.94 | |
粉椴 | Tilia oliveri | 0.92 | |
其他 | Others | 10.12 | 9.03 |
表2 宝天曼国家级自然保护区1 ha落叶阔叶林样地重要值前20位物种的变化(2009-2014年)
Table 2 Importance value (IV) dynamics of top 20 species between 2009 and 2014 in a 1-ha deciduous broad-leaved forest plot in Baotianman National Nature Reserve
物种 Species | 拉丁名 Latin name | 重要值 IV | |
---|---|---|---|
2009 | 2014 | ||
槲栎 | Quercus aliena | 20.46 | 21.65 |
水榆花楸 | Sorbus alnifolia | 16.69 | 19.17 |
三桠乌药 | Lindera obtusiloba | 11.26 | 7.59 |
青榨槭 | Acer davidii | 9.62 | 10.88 |
野茉莉 | Styrax japonicus | 4.94 | 5.72 |
毛樱桃 | Cerasus tomentosa | 3.49 | 3.53 |
连翘 | Forsythia suspensa | 2.49 | 2.75 |
白檀 | Symplocos paniculata | 2.42 | 1.88 |
卫矛 | Euonymus alatus | 2.42 | 2.41 |
秦岭木姜子 | Litsea tsinlingensis | 2.38 | |
元宝槭 | Acer truncatum | 2.24 | 2.70 |
千金榆 | Carpinus cordata | 1.75 | 2.00 |
山梅花 | Philadelphus incanus | 1.59 | 1.35 |
暖木 | Meliosma veitchioru | 1.52 | 1.74 |
泡花树 | Meliosma cuneifolia | 1.44 | 1.59 |
马鞍树 | Maackia hupehensi | 1.55 | |
桦叶荚蒾 | Viburnum betulifolium | 1.32 | 1.59 |
四照花 | Dendrobenthamia japonica var. chinensis | 1.25 | 1.55 |
华东椴 | Tilia japonica | 1.06 | |
老鸹铃 | Styrax hemsleyanus | 1.02 | |
灯台树 | Bothrocaryum controversum | 0.94 | |
粉椴 | Tilia oliveri | 0.92 | |
其他 | Others | 10.12 | 9.03 |
种名 Species | 参数 Parameter | 邻体距离 Neighbourhood distance | ||
---|---|---|---|---|
5 m | 7.5 m | 10 m | ||
槲栎 Quercus aliena | CON | 0 | 0 | 0 |
HET | + | + | 0 | |
CONBA | +++ | + | 0 | |
HETBA | - - - | - - - | - - - | |
水榆花楸 Sorbus alnifolia | CON | +++ | +++ | +++ |
HET | - - - | - - - | - - - | |
CONBA | - - - | - - - | - - - | |
HETBA | +++ | +++ | +++ | |
青榨槭 Acer davidii | CON | 0 | 0 | 0 |
HET | 0 | 0 | 0 | |
CONBA | - - - | - - - | - - - | |
HETBA | 0 | 0 | 0 | |
三桠乌药 Lindera obtusiloba | CON | 0 | + | ++ |
HET | - | - | - | |
CONBA | - - | - - | - - | |
HETBA | 0 | 0 | 0 | |
野茉莉 Styrax japonicus | CON | - - - | - - - | - - - |
HET | 0 | 0 | 0 | |
CONBA | - | - | - | |
HETBA | + | 0 | 0 | |
元宝槭 Acer truncatum | CON | 0 | 0 | 0 |
HET | - - | - | - - | |
CONBA | 0 | - | - - | |
HETBA | 0 | 0 | 0 | |
毛樱桃 Cerasus tomentosa | CON | - - | - - | - |
HET | 0 | 0 | 0 | |
CONBA | 0 | 0 | - | |
HETBA | 0 | 0 | 0 | |
白檀 Symplocos paniculata | CON | 0 | 0 | 0 |
HET | 0 | 0 | 0 | |
CONBA | - | - - | - - | |
HETBA | 0 | 0 | 0 | |
秦岭木姜子 Litsea tsinlingensis | CON | 0 | 0 | 0 |
HET | 0 | 0 | 0 | |
CONBA | 0 | 0 | 0 | |
HETBA | ++ | + | 0 | |
千金榆 Carpinus cordata | CON | 0 | - | - - |
HET | 0 | 0 | 0 | |
CONBA | 0 | 0 | 0 | |
HETBA | 0 | 0 | 0 | |
暖木 Meliosma veitchiorum | CON | - - | - - - | - - |
HET | 0 | 0 | 0 | |
CONBA | 0 | 0 | 0 | |
HETBA | 0 | 0 | + |
表3 宝天曼国家级自然保护区1 ha落叶阔叶林样地不同尺度下的邻体分析结果
Table 3 Result of neighborhood analysis for different scales with different species in a 1-ha deciduous broad-leaved forest plot in Baotianman National Nature Reserve
种名 Species | 参数 Parameter | 邻体距离 Neighbourhood distance | ||
---|---|---|---|---|
5 m | 7.5 m | 10 m | ||
槲栎 Quercus aliena | CON | 0 | 0 | 0 |
HET | + | + | 0 | |
CONBA | +++ | + | 0 | |
HETBA | - - - | - - - | - - - | |
水榆花楸 Sorbus alnifolia | CON | +++ | +++ | +++ |
HET | - - - | - - - | - - - | |
CONBA | - - - | - - - | - - - | |
HETBA | +++ | +++ | +++ | |
青榨槭 Acer davidii | CON | 0 | 0 | 0 |
HET | 0 | 0 | 0 | |
CONBA | - - - | - - - | - - - | |
HETBA | 0 | 0 | 0 | |
三桠乌药 Lindera obtusiloba | CON | 0 | + | ++ |
HET | - | - | - | |
CONBA | - - | - - | - - | |
HETBA | 0 | 0 | 0 | |
野茉莉 Styrax japonicus | CON | - - - | - - - | - - - |
HET | 0 | 0 | 0 | |
CONBA | - | - | - | |
HETBA | + | 0 | 0 | |
元宝槭 Acer truncatum | CON | 0 | 0 | 0 |
HET | - - | - | - - | |
CONBA | 0 | - | - - | |
HETBA | 0 | 0 | 0 | |
毛樱桃 Cerasus tomentosa | CON | - - | - - | - |
HET | 0 | 0 | 0 | |
CONBA | 0 | 0 | - | |
HETBA | 0 | 0 | 0 | |
白檀 Symplocos paniculata | CON | 0 | 0 | 0 |
HET | 0 | 0 | 0 | |
CONBA | - | - - | - - | |
HETBA | 0 | 0 | 0 | |
秦岭木姜子 Litsea tsinlingensis | CON | 0 | 0 | 0 |
HET | 0 | 0 | 0 | |
CONBA | 0 | 0 | 0 | |
HETBA | ++ | + | 0 | |
千金榆 Carpinus cordata | CON | 0 | - | - - |
HET | 0 | 0 | 0 | |
CONBA | 0 | 0 | 0 | |
HETBA | 0 | 0 | 0 | |
暖木 Meliosma veitchiorum | CON | - - | - - - | - - |
HET | 0 | 0 | 0 | |
CONBA | 0 | 0 | 0 | |
HETBA | 0 | 0 | + |
模型类型 Model type | 邻体距离 Neighbourhood distance | |||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|
5 m | 7.5 m | 10 m | ||||||||||
小 Small | 中 Medium | 大 Large | 小 Small | 中 Medium | 大 Large | 小 Small | 中 Medium | 大 Large | ||||
基本模型 Basic model | d+n+h+(sp+20s)random | 1,484.06 | 355.5304 | 380.3177 | 1,483.798 | 355.0877 | 372.4036 | 1,484.64 | 359.2236 | 373.3339 | ||
d+c+b+(sp+20s)random | 1,480.921 | 355.6752 | 378.72 | 1,482.687 | 355.3472 | 371.565 | 1,479.719 | 355.676 | 371.266 | |||
d+n+h+c+b+(sp+20s)random | 1,483.84 | 359.2017 | 381.6294 | 1,485.789 | 358.0195 | 370.1479 | 1,483.541 | 357.485 | 369.087 | |||
d+n+h+c+b+i+(sp+20s)random | 1,485.83 | 356.6697 | 379.306 | 1,487.509 | 356.979 | 372.125 | 1,485.34 | 357.0264 | 370.084 | |||
交互作用 模型 Interaction model | d*(n+h) +(sp+20s)random | 1,484.249 | 359.3985 | 382.5019 | 1,485.738 | 358.9014 | 374.9681 | 1,486.79 | 358.066 | 375.3167 | ||
d*(n+h+i) +(sp+20s)random | 1,488.12 | 359.3586 | 384.9341 | 1,489.323 | 362.611 | 378.8982 | 1,489.957 | 361.7479 | 378.8444 | |||
d*(c+b) +(sp+20s)random | 1,475.82 | 358.6642 | 378.6484 | 1,479.193 | 357.8086 | 375.5305 | 1,481.48 | 359.2018 | 374.0463 | |||
d*(c+b+i) +(sp+20s)random | 1,479.6 | 358.9814 | 380.8261 | 1,473.86 | 361.5722 | 379.2916 | 1,481.13 | 363.1487 | 377.7625 | |||
d*(n+h+c+b) +(sp+20s)random | 1,480.32 | 366.2005 | 383.7345 | 1,475.386 | 365.396 | 377.4086 | 1,479.243 | 375.7429 | 375.7429 | |||
d*(n+h+c+b+i) +(sp+20s)random | 1,474.3 | 366.2815 | 386.4617 | 1,474.083 | 369.1937 | 381.3527 | 1,480.58 | 368.7947 | 379.6699 |
表4 不同尺度下不同广义线性混合模型(GLMMs)的AIC值
Table 4 AIC values of different GLMMs (generalized linear mixed models) at three scales (r) in a 1-ha deciduous broad-leaved forest plot in Baotianman National Nature Reserve
模型类型 Model type | 邻体距离 Neighbourhood distance | |||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|
5 m | 7.5 m | 10 m | ||||||||||
小 Small | 中 Medium | 大 Large | 小 Small | 中 Medium | 大 Large | 小 Small | 中 Medium | 大 Large | ||||
基本模型 Basic model | d+n+h+(sp+20s)random | 1,484.06 | 355.5304 | 380.3177 | 1,483.798 | 355.0877 | 372.4036 | 1,484.64 | 359.2236 | 373.3339 | ||
d+c+b+(sp+20s)random | 1,480.921 | 355.6752 | 378.72 | 1,482.687 | 355.3472 | 371.565 | 1,479.719 | 355.676 | 371.266 | |||
d+n+h+c+b+(sp+20s)random | 1,483.84 | 359.2017 | 381.6294 | 1,485.789 | 358.0195 | 370.1479 | 1,483.541 | 357.485 | 369.087 | |||
d+n+h+c+b+i+(sp+20s)random | 1,485.83 | 356.6697 | 379.306 | 1,487.509 | 356.979 | 372.125 | 1,485.34 | 357.0264 | 370.084 | |||
交互作用 模型 Interaction model | d*(n+h) +(sp+20s)random | 1,484.249 | 359.3985 | 382.5019 | 1,485.738 | 358.9014 | 374.9681 | 1,486.79 | 358.066 | 375.3167 | ||
d*(n+h+i) +(sp+20s)random | 1,488.12 | 359.3586 | 384.9341 | 1,489.323 | 362.611 | 378.8982 | 1,489.957 | 361.7479 | 378.8444 | |||
d*(c+b) +(sp+20s)random | 1,475.82 | 358.6642 | 378.6484 | 1,479.193 | 357.8086 | 375.5305 | 1,481.48 | 359.2018 | 374.0463 | |||
d*(c+b+i) +(sp+20s)random | 1,479.6 | 358.9814 | 380.8261 | 1,473.86 | 361.5722 | 379.2916 | 1,481.13 | 363.1487 | 377.7625 | |||
d*(n+h+c+b) +(sp+20s)random | 1,480.32 | 366.2005 | 383.7345 | 1,475.386 | 365.396 | 377.4086 | 1,479.243 | 375.7429 | 375.7429 | |||
d*(n+h+c+b+i) +(sp+20s)random | 1,474.3 | 366.2815 | 386.4617 | 1,474.083 | 369.1937 | 381.3527 | 1,480.58 | 368.7947 | 379.6699 |
图2 邻域内同种个体数(CON)、异种个体数(HET)、邻域关联指数(NRI)对目标个体存活的影响 small、medium、large分别表示小径级(1 cm ≤ DBH < 5 cm)、中径级(5 cm ≤ DBH < 10 cm)和大径级(DBH ≥ 10 cm); 5 m、7.5 m、10 m分别为不同邻域尺度; ○表示相关不显著(P > 0.05),●、■分别表示在P < 0.05、P < 0.01水平上显著相关。
Fig. 2 Neighbourhood effects of conspecific neighburs (CON), heterospecific neighburs (HET) and neighborhood relatedness index (NRI) on individual survivalSmall, medium and large indicate small DBH class (1 cm ≤ DBH < 5 cm), medium DBH class (5 cm ≤ DBH < 10 cm) and large DBH class (DBH ≥ 10 cm), respectively. The 5 m, 7.5 m and 10 m indicate different neighbourhood scales. ○, ●, ■ indicate correlations at P > 0.05, P < 0.05 and P < 0.01 level, respectively.
图3 不同尺度下不同径级树木个体的邻体效应对其存活的影响 A、B、C分别表示小径级((1 cm ≤ DBH < 5 cm)、中径级(5cm ≤ DBH < 10 cm)和大径级(10 cm ≤ DBH ), 数字1、2、3分别表示5 m、7.5 m、10 m的邻域尺度。○为相关不显著, ●、▲、■分别表示在P < 0.05、P < 0.01、P < 0.001水平上显著相关。
Fig. 3 Neighbourhood effects of individual trees with different DBH size and different scales on survivalA, B and C mean different size with small (1 cm ≤ DBH < 5 cm), medium (5 cm ≤ DBH < 10 cm) and large (DBH ≥ 10 cm) DBH, respectively. The number 1, 2, 3 indicate different neighbourhood scales of 5 m, 7.5 m and 10 m. ○ mean no significance correlation (P > 0.05), and ●、▲、■ indicate different significant correlations at p-vale as 0.05, 0.01 and 0.001, respectively.
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