吉林蛟河针阔混交林乔木幼苗组成及其密度格局影响因素

doi:10.17521/cjpe.2018.0025

摘要/Abstract

摘要：

以吉林蛟河次生针阔混交林42 hm ²固定监测样地中209个幼苗监测站内的乔木幼苗为研究对象, 基于2016和2017年幼苗调查数据, 探究幼苗物种组成、数量动态等特征, 并运用广义线性混合效应模型分析了幼苗密度与生物邻体及其生境因素的相关性。结果表明: (1)该样地内所有幼苗监测站共调查到幼苗4 245株, 分属10科12属18种, 新生幼苗的数量在物种和调查年份间均有明显差异, 其中水曲柳(Fraxinus mandschurica)和紫椴(Tilia amurensis)幼苗表现出大量出生和死亡的动态特征。(2)群落水平上, 幼苗密度与局域同种成体胸高断面积之和、土壤全磷和有效钾等养分含量显著正相关, 适宜大树生长的生境同样适宜幼苗的生长。(3)水曲柳幼苗密度的影响因素与群落水平一致, 红松(Pinus koraiensis)存在明显的生境偏好, 在湿度较小、土壤全磷、有效氮和有效磷含量较低的生境中密度更高。相对于多年生幼苗, 同种成年邻体对当年生幼苗密度影响更加显著。该研究证实了扩散限制和生境过滤共同影响幼苗密度格局, 生物邻体和生境异质性的相对重要性随幼苗物种种类和年龄级变化。

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

Abstract:

Aims Our aim is to explore the composition and temporal dynamics of tree seedlings and to further understand the impacts and relative importance of biotic neighborhood and habitat heterogeneity on seedling density patterns in a secondary mixed conifer and broad-leaved forest in Jiaohe, Jilin Province, North-eastern China.

Methods We established 209 seedling census stations in the mixed conifer and broad-leaved forest. Based on two seedling censuses conducted in 2016 and 2017, we analyzed species composition and quantitative dynamics of seedlings. Generalized mixed linear models were used to test the relative effects of biotic neighborhood and habitat heterogeneity on seedling density patterns.

Important findings (1) A total of 4 245 seedlings were investigated in all seedling census stations of this plot, belonging to 12 genera, 10 families and 18 species. The number of new-born seedlings varied greatly among species and census years. Fraxinus mandschurica and Tilia amurensis seedlings occurred and died in large numbers. (2) At the community level, the sum of conspecific adult basal area, the soil total phosphorus and available potassium exhibited significant positive effects on seedling density. The habitat suitable for trees was also suitable for seedlings. (3) The factors influencing seedling density of Fraxinus mandschurica were consistent with those of seedling density at the community level. Pinus koraiensis had significant habitat preference, it had higher density in the habitat with lower moisture, soil total phosphorus, available nitrogen, and available phosphorus. Conspecific adult neighbor effects on annual seedling density was more significant than on perennial seedling density. This study suggests that dispersal limitation and habitat filtering together affect seedling density pattern, and the relative importance of biotic neighbors and habitat heterogeneity varies among seedling species and age classes.

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

孟令君, 姚杰, 秦江环, 范春雨, 张春雨, 赵秀海. 吉林蛟河针阔混交林乔木幼苗组成及其密度格局影响因素. 植物生态学报, 2018, 42(6): 653-662. DOI: 10.17521/cjpe.2018.0025

MENG Ling-Jun, YAO Jie, QIN Jiang-Huan, FAN Chun-Yu, ZHANG Chun-Yu, ZHAO Xiu-Hai. Drivers of composition and density pattern of tree seedlings in a secondary mixed conifer and broad-leaved forest, Jiaohe, Jilin, China. Chinese Journal of Plant Ecology, 2018, 42(6): 653-662. DOI: 10.17521/cjpe.2018.0025

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https://www.plant-ecology.com/CN/Y2018/V42/I6/653

图/表 10

参考文献 52

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土壤变量 Soil variables	PC1	PC2	PC3
全氮 Total nitrogen	0.320	-0.337	0.538
全磷 Total phosphorus	0.450	0.182	-0.218
全钾 Total potassium	-0.371	-0.492	-
有机质 Organic matter	0.378	-0.523	0.233
有效氮 Available nitrogen	0.421	-0.319	-0.359
有效磷 Available phosphorus	-	-0.258	-0.688
有效钾 Available potassium	0.480	0.411	-
变异解释 Proportion of variance	34.224%	18.990%	16.428%

土壤变量 Soil variables	PC1	PC2	PC3
全氮 Total nitrogen	0.320	-0.337	0.538
全磷 Total phosphorus	0.450	0.182	-0.218
全钾 Total potassium	-0.371	-0.492	-
有机质 Organic matter	0.378	-0.523	0.233
有效氮 Available nitrogen	0.421	-0.319	-0.359
有效磷 Available phosphorus	-	-0.258	-0.688
有效钾 Available potassium	0.480	0.411	-
变异解释 Proportion of variance	34.224%	18.990%	16.428%

树种 Species	幼苗数量 No. of seedlings		新生幼苗数量(新生率) No. of recruitment (Recruitment rate (%))		死亡幼苗数量(死亡率) No. of mortality (Mortality rate (%))	幼苗密度 Seedling density (Ind.·m^-2)		分布的样方数 No. of stations occupied		重要值 Important value
树种 Species	2016	2017	2016	2017	2016-2017	2016	2017	2016	2017	2016	2017
水曲柳 Fraxinus mandschurica	1 769	796	871 (49.2)	213 (26.8)	1 186 (67.0)	2.116	0.952	498	317	52.28	42.11
色木槭 Acer mono	199	177	7 (3.5)	16 (9.0)	38 (19.1)	0.238	0.212	143	124	9.59	12.45
紫椴 Tilia amurensis	306	124	274 (89.5)	89 (71.8)	271 (88.6)	0.366	0.148	169	81	12.58	8.38
东北枫 Acer mandshuricum	127	159	12 (9.4)	44 (27.7)	12 (9.4)	0.152	0.19	100	120	6.5	11.68
红松 Pinus koraiensis	192	121	156 (81.3)	35 (28.9)	106 (55.2)	0.230	0.145	94	60	7.38	7.08
簇毛枫 Acer barbinerve	101	107	17 (16.8)	14 (13.1)	8 (7.9)	0.121	0.128	34	31	3.22	4.99
胡桃楸 Juglans mandshurica	43	29	20 (46.5)	14 (48.3)	28 (65.1)	0.051	0.035	40	25	2.46	2.31
杉松 Abies holophylla	35	29	13 (37.1)	5 (17.2)	11 (31.4)	0.042	0.035	28	24	1.81	2.25
蒙古栎 Quercus mongolica	1	62	0 (0)	62 (100)	1 (100)	0.001	0.074	1	27	0.06	3.41
千金榆 Carpinus cordata	30	21	14 (46.7)	6 (28.6)	15 (50)	0.036	0.025	26	19	1.64	1.72
裂叶榆 Ulmus laciniata	24	21	0 (0)	0 (0)	3 (12.5)	0.029	0.025	16	17	1.10	1.61
糠椴 Tilia mandshurica	10	6	4 (40.0)	1 (16.7)	5 (50)	0.012	0.007	8	5	0.52	0.47
春榆 Ulmus davidiana var. japonica	9	5	3 (33.3)	1 (20.0)	5 (55.6)	0.011	0.006	8	4	0.50	0.38
三花枫 Acer triflorum	2	4	1 (50)	2 (50)	0	0.002	0.005	2	4	0.12	0.35
黄檗 Phellodendron amurense	2	3	1 (50.0)	3 (100)	2 (100)	0.002	0.004	2	3	0.12	0.26
青楷枫 Acer tegmentosum	2	3	0 (0)	3 (100)	2 (100)	0.002	0.004	2	2	0.12	0.21
毛榛 Corylus mandshurica	0	4	0	4 (100)	0	0	0.005	0	2	0	0.24
水榆花楸 Sorbus alnifolia	0	1	0	1 (100)	0	0	0.001	0	1	0	0.09
共计 Total	2 852	1 672	1 393 (48.8)	513 (30.7)	1 693 (59.4)	3.411	2.001

树种 Species	幼苗数量 No. of seedlings		新生幼苗数量(新生率) No. of recruitment (Recruitment rate (%))		死亡幼苗数量(死亡率) No. of mortality (Mortality rate (%))	幼苗密度 Seedling density (Ind.·m^-2)		分布的样方数 No. of stations occupied		重要值 Important value
树种 Species	2016	2017	2016	2017	2016-2017	2016	2017	2016	2017	2016	2017
水曲柳 Fraxinus mandschurica	1 769	796	871 (49.2)	213 (26.8)	1 186 (67.0)	2.116	0.952	498	317	52.28	42.11
色木槭 Acer mono	199	177	7 (3.5)	16 (9.0)	38 (19.1)	0.238	0.212	143	124	9.59	12.45
紫椴 Tilia amurensis	306	124	274 (89.5)	89 (71.8)	271 (88.6)	0.366	0.148	169	81	12.58	8.38
东北枫 Acer mandshuricum	127	159	12 (9.4)	44 (27.7)	12 (9.4)	0.152	0.19	100	120	6.5	11.68
红松 Pinus koraiensis	192	121	156 (81.3)	35 (28.9)	106 (55.2)	0.230	0.145	94	60	7.38	7.08
簇毛枫 Acer barbinerve	101	107	17 (16.8)	14 (13.1)	8 (7.9)	0.121	0.128	34	31	3.22	4.99
胡桃楸 Juglans mandshurica	43	29	20 (46.5)	14 (48.3)	28 (65.1)	0.051	0.035	40	25	2.46	2.31
杉松 Abies holophylla	35	29	13 (37.1)	5 (17.2)	11 (31.4)	0.042	0.035	28	24	1.81	2.25
蒙古栎 Quercus mongolica	1	62	0 (0)	62 (100)	1 (100)	0.001	0.074	1	27	0.06	3.41
千金榆 Carpinus cordata	30	21	14 (46.7)	6 (28.6)	15 (50)	0.036	0.025	26	19	1.64	1.72
裂叶榆 Ulmus laciniata	24	21	0 (0)	0 (0)	3 (12.5)	0.029	0.025	16	17	1.10	1.61
糠椴 Tilia mandshurica	10	6	4 (40.0)	1 (16.7)	5 (50)	0.012	0.007	8	5	0.52	0.47
春榆 Ulmus davidiana var. japonica	9	5	3 (33.3)	1 (20.0)	5 (55.6)	0.011	0.006	8	4	0.50	0.38
三花枫 Acer triflorum	2	4	1 (50)	2 (50)	0	0.002	0.005	2	4	0.12	0.35
黄檗 Phellodendron amurense	2	3	1 (50.0)	3 (100)	2 (100)	0.002	0.004	2	3	0.12	0.26
青楷枫 Acer tegmentosum	2	3	0 (0)	3 (100)	2 (100)	0.002	0.004	2	2	0.12	0.21
毛榛 Corylus mandshurica	0	4	0	4 (100)	0	0	0.005	0	2	0	0.24
水榆花楸 Sorbus alnifolia	0	1	0	1 (100)	0	0	0.001	0	1	0	0.09
共计 Total	2 852	1 672	1 393 (48.8)	513 (30.7)	1 693 (59.4)	3.411	2.001

变量 Variables	数据 Data
变量 Variables	最小值 Min.		最大值 Max.		平均值 Mean
生物邻体变量 Biotic neighbors variables
同种成体胸高断面积 Conspecific adult basal area (ConBa)		0		2.466 1	0.407 2
异种成体胸高断面积 Heterospecific adult basal area (HetBa)		1.233		8.151	3.56
生境变量 Habitat variables
林冠开阔度 Canopy openness		0.989		3.982	2.148
土壤含水量 Moisture (%)		13		76.4	26.74
土壤pH值 Soil pH value		3.96		5.44	4.603
第一主成分 The first principal component (PC1)		-3.140 6		4.209 1	0.188 2
第二主成分 The second principal component (PC2)		-6.833 9		3.225 3	0.312 3
第三主成分 The third principal component (PC3)		-4.091 0		2.157 4	0.176 6