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

doi:10.17520/biods.2015134

生物多样性 ›› 2016, Vol. 24 ›› Issue (6): 639-657. DOI: 10.17520/biods.2015134

• 研究报告: 植物多样性 • 上一篇下一篇

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

刘晓静¹, 任思远², 李鹿鑫², 叶永忠², 袁志良², 王婷^2,,A;^*()

1 河南宝天曼国家级自然保护区管理局, 河南内乡 474350
2 河南农业大学, 郑州 450002

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:
国家自然科学基金(31270493)、中国科学院植物研究所植被与环境变化国家重点实验室2012 年开放课题(LVEC-2012kf06)和林业公益性行业科研专项“气候变化对森林水碳平衡影响及适应性生态恢复(201404201)”

Detecting density dependence on tree survival in a deciduous broad- leaved forest in Baotianman National Nature Reserve

Xiaojing Liu¹, Siyuan Ren², Luxin Li², Yongzhong Ye², Zhiliang Yuan², Ting Wang^2,^*()

1 Baotianman National Nature Reserve Administrative Bureau, Neixiang, Henan 474350
2 Henan Agricultural University, Zhengzhou 450002

Received:2016-02-15 Accepted:2016-06-06 Online:2016-06-20 Published:2016-06-20
Contact: Wang Ting

摘要/Abstract

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。

关键词: 密度制约, 广义线性混合模型, 邻体分析, 谱系密度制约

Abstract

It has been found that density dependence plays a crucial role in determination of species distribution and coexistence, and the effect of negative density dependence also plays an important part among phylogenetically related species. Based on two census datasets in 2009 and 2014 of the 1-ha deciduous broad- leaved forest plot in Baotianman National Nature Reserve, generalized linear mixed models (GLMMs) were used to examine the density dependence on individual survival of the top 11 species (with high important value) at different neighborhood scales and different DBH sizes (1 cm ≤ DBH< 5 cm, 5 cm ≤ DBH < 10 cm, DBH ≥ 10 cm). The results showed that (1) mean annual mortality and recruitment rate of individuals (DBH ≥ 1 cm) were 5.85% and 0.27%, respectively; (2) survival rate of 5 tree species was negatively correlated with abundance or neighbor conspecific basal area; (3) density dependence and phylogenetic diversity exerted significant effects on survival of small individuals (at 5 m neighborhood scale), whereas no significant influence on medium trees. With increasing tree size, phylogenetic diversity exerted important effect on large individuals at 7.5 m and 10 m neighborhood scales. In conclusion, density dependence and phylogenetic diversity play different roles on tree survival at different life stages and neighborhood scales in the deciduous broad-leaved forest in Baotianman National Nature Reserve.

Key words: density dependence, generalized linear mixed models (GLMMs), neighborhood analysis, phylogenetic density dependence

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, 639-657. 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, 639-657. DOI: 10.17520/biods.2015134.

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链接本文: https://www.biodiversity-science.net/CN/10.17520/biods.2015134

https://www.biodiversity-science.net/CN/Y2016/V24/I6/639

图/表 7

参考文献 35

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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, %)
径级 DBH size	2009	2014	2009	2014	死亡数(死亡率) Number of mortality (mortality rate, %)	新增数(增员率) Number of recruitment (recruitment rate, %)
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

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, %)
径级 DBH size	2009	2014	2009	2014	死亡数(死亡率) Number of mortality (mortality rate, %)	新增数(增员率) Number of recruitment (recruitment rate, %)
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

物种 Species	拉丁名 Latin name	重要值 IV
物种 Species	拉丁名 Latin name	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

物种 Species	拉丁名 Latin name	重要值 IV
物种 Species	拉丁名 Latin name	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
种名 Species	参数 Parameter	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	+

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

Detecting density dependence on tree survival in a deciduous broad- leaved forest in Baotianman National Nature Reserve

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参考文献 35

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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

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