长白山次生针阔混交林乔木幼苗存活的影响因素分析

doi:10.17521/cjpe.2015.0366

摘要/Abstract

摘要：

基于长白山次生针阔混交林样地, 以520个1 m × 1 m幼苗样方中胸径小于1 cm的乔木幼苗为研究对象, 选取2013年和2014年的幼苗调查数据, 运用广义线性混合模型(GLMM)分析了生物邻体和生境异质性对幼苗存活的影响, 探讨了次生针阔混交林幼苗存活影响因素及物种共存机制。结果表明: (1)适宜大树生长的局域生境同样也适宜幼苗的存活, 幼苗存活率与土壤含水量和有效氮等土壤养分显著正相关。(2)幼苗个体之间存在明显的竞争, 较多的幼苗邻体显著降低幼苗的存活率。同种大树邻体和同种幼苗邻体与幼苗存活显著负相关, 表明存在负密度制约效应。(3)随着幼苗年龄的增加, 生境异质性对幼苗存活的影响逐渐增大。该研究证实了密度制约效应和生境异质性对幼苗存活有着重要影响, 其相对重要性随着幼苗年龄级、功能群以及物种种类而变化。

关键词: 次生针阔混交林, 广义线性混合模型, 乔木幼苗存活, 生物邻体, 生境异质性

Abstract:

Aims Our study aimed to understand the effects and the relative importance of biotic neighborhood and habitat heterogeneity for tree seedling survival in a secondary mixed conifer and broad-leaved forest in Changbai Mountain, north-eastern China.
Methods The generalized mixed linear model was used to examine the relative effects of biotic neighborhood and habitat heterogeneity on seedling survival over two years.
Important findings Our results showed that both biotic neighborhood and habitat heterogeneity had significant effects on the seedling survival at community level. The local environment suitable for the adult growth was also suitable for seedling survival. The soil moisture and soil available nitrogen exhibited significant positive effects on seedling survival. On the other hand, seedling density had significant negative effects on seedling survival due to the individual competition. Particularly, we found significant negative density-dependent effects on seedling survival which was caused by conspecific adult and seedling neighbors. As expected, with the increasing of seedling survival age, the habitat heterogeneity became more important on seedling survival. These results suggest that both local biotic neighborhood and habitat heterogeneity drive seedling survival in this temperate forest, and their relative importance varies with different seedling age classes and species traits.

Key words: secondary mixed conifer and broad-leaved forest, generalized linear mixed model, tree seedling survival, biotic neighborhood, habitat heterogeneity

刘帅, 廖嘉星, 肖翠, 范秀华. 长白山次生针阔混交林乔木幼苗存活的影响因素分析. 植物生态学报, 2016, 40(7): 711-722. DOI: 10.17521/cjpe.2015.0366

Shuai LIU, Jia-Xing LIAO, Cui XIAO, Xiu-Hua FAN. Effects of biotic neighbors and habitat heterogeneity on tree seedling survival in a secondary mixed conifer and broad-leaved forest in Changbai Mountain. Chinese Journal of Plant Ecology, 2016, 40(7): 711-722. DOI: 10.17521/cjpe.2015.0366

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2015.0366

https://www.plant-ecology.com/CN/Y2016/V40/I7/711

图/表 9

图1 幼苗样方在样地中的分布。□, 种子收集器; ■, 幼苗样方。

Fig. 1 Distribution of seedling quadrats within the plot. □, seed trap; ■, seedling quadrats.

表1 土壤变量分别在3个主成成分上的载荷值

Table 1 Soil variables loadings on the three PCAs

土壤变量 Soil variable	PCA1	PCA2	PCA3
有机质 Organic matter	-0.455	-	0.367
全磷 Total phosphorus	-0.476	-	-0.202
有效磷 Available phosphorus	-0.223	0.721	-0.372
全钾 Total potassium	0.395	0.166	-0.353
有效钾 Available potassium	-0.435	0.344	-
全氮 Total nitrogen	-0.396	-0.285	-
有效氮 Available nitrogen	-0.117	-0.489	-0.745
变量解释 Variation explained	39.9%	15.6%	15.1%

表2 幼苗存活模型中出现的参数

Table 2 Parameters present in models of seedling survival

变量 Variables	数据 Data
变量 Variables	最小值 Min	最大值 Max	平均值 Mean	中值 Median
幼苗高度 Seedling height (h, cm)	1	98	11.82	7
大树邻体作用 Adult effect
同种个体数 No. of conspecific individuals (ind.c)	0	76	8.69	5.00
异种个体数 No. of heterospecific individuals (ind.h)	19	132	73.34	73.50
同种胸高断面积 Conspecific basal area (ba.c)	0.000	0.510	0.007	0.001
异种胸高断面积 Heterospecific basal area (ba.h)	0.002	4.700	0.096	0.047
幼苗邻体作用 Seedling effect
同种幼苗 Conspecific seedlings (s.c)	1	20	4.39	3
异种幼苗 Heterospecific seedlings (s.h)	0	37	6.89	5
幼苗年龄 Seedling age	1	未知 Unknown	-	-
土壤特性 Soil property
第一主成分 The first principal component (PC1)	-3.901	3.646	0.572	0.789
第二主成分 The second principal component (PC2)	-1.978	2.840	-0.112	-0.144
第三主成分 The third principal component (PC3)	-2.286	4.385	0.186	0.169
土壤pH值 Soil pH value	4.29	6.73	5.22	5.18
土壤含水量 Soil water content (%)	29.06	69.46	40.31	39.78
林冠开阔度 Canopy openness (%)	6.76	29.73	11.96	11.88

图2 群落水平乔木幼苗存活最优模型各参数的优势比率。圆圈表示各参数在模型中的估计值, 水平线表示95%的置信区间, 显著的参数估计值用实心圆表示, 参数缩写见表2。

Fig. 2 Odds. ratios for model parameters of tree seedling survival for the fullest most likely models at community level. Circles show estimated Odds. ratios for each parameter, with 95% confidence limits indicated by horizontal lines. Odds. ratios significantly different from 1 (95% confidence limits do not overlap 1) are indicated by filled circles. See Table 2 for variable abbreviations.

表3 幼苗存活模型的赤池信息量准则(AIC)和ΔAIC值

Table 3 Akaike’s information criterion (AIC) and ΔAIC values of models of seedling survival

数据子集 Data subsets	备选模型 Candidate model
	零模型 Null model		生物模型 Biotic model		生境模型 Habitat model		生物生境模型 Biotic + Habitat model
	AIC	ΔAIC	AIC	ΔAIC	AIC	ΔAIC	AIC	ΔAIC
群落水平 Community level	1 683.30	361.94	1 563.96	242.61	1 346.31	24.96	1 321.36	0.00
种子传播方式 Seed dispersal mode
风媒传播 Wind dispersal	788.51	67.56	792.04	71.09	724.23	3.28	720.95	0.00
重力传播 Gravity dispersal	253.48	6.79	289.26	42.57	246.69	0.00	249.40	2.71
年龄级 Age class
一年生 One year old	994.63	257.96	882.44	145.78	766.17	29.50	736.66	0.00
二至三年生 Two to three years old	266.32	35.69	258.77	28.14	230.63	0.00	235.21	4.58
四年生以上 Above four years old	290.00	9.53	297.74	17.27	280.47	0.00	287.74	7.28
耐阴性 Shade tolerance
阳性 Light-demanding	344.57	0.72	368.37	24.51	343.85	0.00	353.90	10.05
中性 Mid shade-tolerant	108.20	0.00	117.34	9.14	113.90	5.70	113.70	5.50
阴性 Shade-tolerant	587.24	40.95	585.02	38.73	547.34	1.04	546.30	0.00
物种 Species
紫椴 Tilia amurensis	209.68	4.31	218.52	13.15	205.37	0.00	206.29	0.92
簇毛槭 Acer barbinerve	255.25	56.51	236.21	37.47	207.61	8.87	198.74	0.00

图3 不同年龄级乔木幼苗存活最优模型各参数的优势比率, 圆圈表示各参数在模型中的估计值, 水平线表示95%的置信区间, 显著的参数估计值用实心圆表示, 参数缩写见表2。

Fig. 3 Odds. ratios for model parameters of tree seedling survival of different age classes for the fullest most likely models. Circles show estimated Odds. ratios for each parameter, with 95% confidence limits indicated by horizontal lines. Odds. ratios significantly different from 1 (95% confidence limits do not overlap 1) are indicated by filled circles. See Table 2 for variable abbreviations.

图4 不同种子传播模式乔木幼苗存活最优模型各参数的优势比率, 圆圈表示各参数在模型中的估计值, 水平线表示95%的置信区间, 显著的参数估计值用实心圆表示, 参数缩写见表2。

Fig. 4 Odds. ratios for model parameters of tree seedling survival of dispersal-mode groups for the fullest most likely models in Table 3, Circles show estimated Odds. ratios for each parameter, with 95% confidence limits (CL) indicated by horizontal lines. Odds. ratios significantly different from 1 (95% CL do not overlap 1) are indicated by filled circles. See Table 2 for variable abbreviations.

图5 不同耐阴性乔木幼苗存活最优模型各参数的优势比率, 圆圈表示各参数在模型中的估计值, 水平线表示95%的置信区间, 显著的参数估计值用实心圆表示, 参数缩写见表2。

Fig. 5 Odds. ratios for model parameters of tree seedling survival of shade tolerant-mode groups for the fullest most likely models , Circles show estimated Odds. ratios for each parameter, with 95% confidence limits (CL) indicated by horizontal lines. Odds. ratios significantly different from 1 (95% CL do not overlap 1) are indicated by filled circles. See Table 2 for variable abbreviations.

图6 不同物种乔木幼苗存活最优模型各参数的优势比率, 圆圈表示各参数在模型中的估计值, 水平线表示95%的置信区间, 显著的参数估计值用实心圆表示, 参数缩写见表2。

Fig. 6 Odds. ratios for model parameters of tree seedling survival of different species for the fullest most likely models, circles show estimated Odds. ratios for each parameter, with 95% confidence limits (CL) indicated by horizontal lines. Odds. ratios significantly different from 1 (95% CL do not overlap 1) are indicated by filled circles. See Table 2 for variable abbreviations..

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