阔叶红松林不同演替阶段灌木叶片碳氮磷化学计量特征及其影响因素

doi:10.17521/cjpe.2021.0101

摘要/Abstract

摘要：

灌木是森林生态系统的重要组成部分, 对于演替进程中灌木叶片化学计量特征的研究, 有助于全面理解和预测森林演替过程。该研究以黑龙江凉水国家自然保护区内处于阔叶红松(Pinus koraiensis)林不同演替阶段中的白桦(Betula platyphylla)次生林、落叶阔叶混交林、针阔混交林、阔叶红松林的灌木为研究对象, 分析其叶片的碳(C)、氮(N)、磷(P)化学计量特征差异, 并利用层次分割方法检验其与土壤、物种多样性的关系。主要结果为: 1)随着演替的进行, 阔叶红松林的叶片N含量显著高于其他3种林型, P含量与白桦次生林无显著差异, 但显著高于其他两种林型; 2)土壤N、P含量与个体尺度上的叶片N含量均呈显著正相关关系, 土壤P含量与叶片P含量呈显著正相关关系; 3)群落尺度上, 物种多样性和土壤化学性质共解释叶片N含量变异的82%和叶片P含量变异的62%; 4)群落尺度上Shannon多样性指数与灌木叶片的N、P含量呈显著正相关关系, 与灌木叶片的C:N、C:P呈显著负相关关系。总之, 阔叶红松林4个演替阶段灌木均受到氮限制; 相较于土壤的化学性质, 物种多样性更好地解释了灌木化学计量的变异。

关键词: 物种多样性, 养分限制, 土壤, 小兴安岭, 阔叶红松林

Abstract:

Aims Shrubs are an important component of forest ecosystems. This study investigated changes in the stoichiometric characteristics of shrub leaves during forest succession in order to understand and predict the processes of forest succession.

Methods The study was conducted in the Liangshui National Nature Reserve of Heilongjiang Province, with forest stands at different successional stages of mixed broadleaved-Korean pine (Pinus koraiensis) forest representing secondary birch (Betula platyphylla) forest, mixed deciduous broad-leaved forest, mixed coniferous and broad-leaved forest, and mixed broadleaved-Korean pine forest. Measurements were made on carbon (C), nitrogen (N) and phosphorus (P) contents in leaves of the understory shrubs and soil, and the stoichiometric characteristics of shrub leaves and relationships with soil stoichiometry were examined with hierarchical analysis.

Important findings The N content in shrub leaves was significantly higher in the mixed broadleaved-Korean pine forest than in other three forest types; the P content was significantly higher in the mixed broadleaved-Korean pine forest than in two other forest types except the secondary birch forest. Soil N and P contents were significantly and positively correlated with leaf N content at individual scale, and soil P concentration was significantly and positively correlated with leaf P content. At the community level, 82% of leaf N content variation and 62% of leaf P content variation were explained by species diversity and soil chemical properties; the Shannon diversity index was significantly and positively correlated with the N and P contents in shrub leaves, and negatively with the leaf C:N ratio and C:P ratio. In conclusion, shrubs in mixed broadleaved-Korean pine forests at the four successional stages were all N-limited, and species diversity better explains the stoichiometric variations in understory shrubs than soil chemical properties.

Key words: species diversity, nutrient limitation, soil, Xiao Hinggan Mountains, mixed broadleaved-Korean pine forest

宋语涵, 张鹏, 金光泽. 阔叶红松林不同演替阶段灌木叶片碳氮磷化学计量特征及其影响因素. 植物生态学报, 2021, 45(9): 952-960. DOI: 10.17521/cjpe.2021.0101

SONG Yu-Han, ZHANG Peng, JIN Guang-Ze. Characteristics of shrub leaf carbon, nitrogen and phosphorus stoichiometry and influencing factors in mixed broadleaved-Korean pine forests at different successional stages. Chinese Journal of Plant Ecology, 2021, 45(9): 952-960. DOI: 10.17521/cjpe.2021.0101

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2021.0101

https://www.plant-ecology.com/CN/Y2021/V45/I9/952

图/表 5

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演替阶段 Successional stage	乔木组成 Tree composition	灌木组成 Shrub composition
演替阶段I (白桦次生林) Successional stage I (Secondary birch forest)	5白桦; 2红松; 1兴安落叶松; 1水曲柳+春榆+色木槭+山杨+黄檗-枫桦-红皮云杉-裂叶榆-臭冷杉-胡桃楸-紫椴 5 Betula platyphylla; 2 Pinus koraiensis; 1 Larix gmelinii; 1 Fraxinus mandschurica + Ulmus davidiana var. japonica + Acer pictum subsp. mono + Populus davidiana + Phellodendron amurense - Betula costata - Picea koraiensis - Ulmus laciniata - Abies nephrolepis - Juglans mandshurica - Tilia amurensis	7暴马丁香; 1东北山梅花; 1刺五加+珍珠梅+早花忍冬+毛榛子-光萼溲疏 7 Syringa reticulata subsp. amurensis; 1 Philadelphus schrenkii; 1 Acanthopanax senticosus + Sorbaria sorbifolia + Lonicera praeflorens + Corylus mandshurica - Deutzia glabrata
演替阶段II (落叶阔叶混交林) Successional stage II (Mixed deciduous broad-leaved forest)	4水曲柳; 2山杨; 1紫椴; 1色木槭; 1白桦+胡桃楸+鱼鳞云杉-春榆-红松-红皮云杉-青楷槭-稠李-裂叶榆-蒙古栎 4 Fraxinus mandschurica; 2 Populus davidiana; 1 Tilia amurensis; 1 Betula platyphylla + Juglans mandshurica + Picea jezoensis var. microsperma - Ulmus laciniata - Pinus koraiensis - Picea koraiensis - Acer tegmentosum - Padus racemose - Ulmus laciniata - Quercus mongolica	7暴马丁香; 2毛榛; 1瘤枝卫矛+东北山梅花+刺五加+光萼溲疏-早花忍冬 7 Syringa reticulata subsp. amurensis; 2 Corylus mandshurica; 1 Euonymus verrucosus + Philadelphus schrenkii + Acanthopanax senticosus + Deutzia glabrata - Lonicera praeflorens
演替阶段III (针阔混交林) Succession stage III (Mixed coniferous and broad-leaved forest)	4臭冷杉; 4红松; 1春榆; 1红皮云杉+鱼鳞云杉+白桦+色木槭-花楷槭-枫桦-紫椴-胡桃楸-稠李 4 Abies nephrolepis; 4 Pinus koraiensis; 1 Ulmus davidiana var. japonica; 1 Picea koraiensis + Picea jezoensis var. microsperma + Betula platyphylla + Acer pictum subsp. mono - Acer ukurunduense - Betula costata - Tilia amurensis - Juglans mandshurica - Padus racemosa	7暴马丁香; 2毛榛+瘤枝卫矛+早花忍冬-刺五加-光萼溲疏-珍珠梅 7 Syringa reticulata subsp. amurensis; 2 Corylus mandshurica + Euonymus verrucosus + Lonicera praeflorens - Acanthopanax senticosus - Deutzia glabrata - Sorbaria sorbifolia
演替阶段IV (阔叶红松林) Succession stage IV (Mixed broadleaved-Korean pine forest)	4红松; 2枫桦; 1色木槭; 1鱼鳞云杉; 1红皮云杉+青楷槭+紫椴+白桦+黄檗+裂叶榆-水曲柳-山杨-稠李-臭冷杉-花楷槭-春榆-胡桃楸 4 Pinus koraiensis; 2 Betula costata; 1 Acer pictum subsp. mono; 1 Picea jezoensis var. microsperma; 1 Picea koraiensis + Acer tegmentosum + Tilia amurensis + Betula platyphylla + Phellodendron amurense + Ulmus laciniata - Fraxinus mandschurica - Populus davidiana - Padus racemosa - Abies nephrolepis - Acer ukurunduense - Ulmus davidiana var. japonica - Juglans mandshurica	2东北山梅花; 2瘤枝卫矛; 1暴马丁香; 1光萼溲疏; 1毛榛; 1早花忍冬; 1刺五加; 1龙牙楤木-绣线菊-东北茶藨子 2 Philadelphus schrenkii; 2 Euonymus verrucosus; 1 Syringa reticulata subsp. amurensis; 1 Deutzia glabrata; 1 Corylus mandshurica; 1 Lonicera praeflorens; 1 Acanthopanax senticosus; 1 Aralia elata - Spiraea salicifolia - Ribes mandshuricum

演替阶段 Successional stage	乔木组成 Tree composition	灌木组成 Shrub composition
演替阶段I (白桦次生林) Successional stage I (Secondary birch forest)	5白桦; 2红松; 1兴安落叶松; 1水曲柳+春榆+色木槭+山杨+黄檗-枫桦-红皮云杉-裂叶榆-臭冷杉-胡桃楸-紫椴 5 Betula platyphylla; 2 Pinus koraiensis; 1 Larix gmelinii; 1 Fraxinus mandschurica + Ulmus davidiana var. japonica + Acer pictum subsp. mono + Populus davidiana + Phellodendron amurense - Betula costata - Picea koraiensis - Ulmus laciniata - Abies nephrolepis - Juglans mandshurica - Tilia amurensis	7暴马丁香; 1东北山梅花; 1刺五加+珍珠梅+早花忍冬+毛榛子-光萼溲疏 7 Syringa reticulata subsp. amurensis; 1 Philadelphus schrenkii; 1 Acanthopanax senticosus + Sorbaria sorbifolia + Lonicera praeflorens + Corylus mandshurica - Deutzia glabrata
演替阶段II (落叶阔叶混交林) Successional stage II (Mixed deciduous broad-leaved forest)	4水曲柳; 2山杨; 1紫椴; 1色木槭; 1白桦+胡桃楸+鱼鳞云杉-春榆-红松-红皮云杉-青楷槭-稠李-裂叶榆-蒙古栎 4 Fraxinus mandschurica; 2 Populus davidiana; 1 Tilia amurensis; 1 Betula platyphylla + Juglans mandshurica + Picea jezoensis var. microsperma - Ulmus laciniata - Pinus koraiensis - Picea koraiensis - Acer tegmentosum - Padus racemose - Ulmus laciniata - Quercus mongolica	7暴马丁香; 2毛榛; 1瘤枝卫矛+东北山梅花+刺五加+光萼溲疏-早花忍冬 7 Syringa reticulata subsp. amurensis; 2 Corylus mandshurica; 1 Euonymus verrucosus + Philadelphus schrenkii + Acanthopanax senticosus + Deutzia glabrata - Lonicera praeflorens
演替阶段III (针阔混交林) Succession stage III (Mixed coniferous and broad-leaved forest)	4臭冷杉; 4红松; 1春榆; 1红皮云杉+鱼鳞云杉+白桦+色木槭-花楷槭-枫桦-紫椴-胡桃楸-稠李 4 Abies nephrolepis; 4 Pinus koraiensis; 1 Ulmus davidiana var. japonica; 1 Picea koraiensis + Picea jezoensis var. microsperma + Betula platyphylla + Acer pictum subsp. mono - Acer ukurunduense - Betula costata - Tilia amurensis - Juglans mandshurica - Padus racemosa	7暴马丁香; 2毛榛+瘤枝卫矛+早花忍冬-刺五加-光萼溲疏-珍珠梅 7 Syringa reticulata subsp. amurensis; 2 Corylus mandshurica + Euonymus verrucosus + Lonicera praeflorens - Acanthopanax senticosus - Deutzia glabrata - Sorbaria sorbifolia
演替阶段IV (阔叶红松林) Succession stage IV (Mixed broadleaved-Korean pine forest)	4红松; 2枫桦; 1色木槭; 1鱼鳞云杉; 1红皮云杉+青楷槭+紫椴+白桦+黄檗+裂叶榆-水曲柳-山杨-稠李-臭冷杉-花楷槭-春榆-胡桃楸 4 Pinus koraiensis; 2 Betula costata; 1 Acer pictum subsp. mono; 1 Picea jezoensis var. microsperma; 1 Picea koraiensis + Acer tegmentosum + Tilia amurensis + Betula platyphylla + Phellodendron amurense + Ulmus laciniata - Fraxinus mandschurica - Populus davidiana - Padus racemosa - Abies nephrolepis - Acer ukurunduense - Ulmus davidiana var. japonica - Juglans mandshurica	2东北山梅花; 2瘤枝卫矛; 1暴马丁香; 1光萼溲疏; 1毛榛; 1早花忍冬; 1刺五加; 1龙牙楤木-绣线菊-东北茶藨子 2 Philadelphus schrenkii; 2 Euonymus verrucosus; 1 Syringa reticulata subsp. amurensis; 1 Deutzia glabrata; 1 Corylus mandshurica; 1 Lonicera praeflorens; 1 Acanthopanax senticosus; 1 Aralia elata - Spiraea salicifolia - Ribes mandshuricum

叶片化学计量 Leaf stoichiometry	R²	物种多样性解释度 Interpretation of species diversity (%)			土壤化学性质解释度 Interpretation of soil chemical properties (%)
叶片化学计量 Leaf stoichiometry	R²	H′	S	J	SC	SN	SP
LC	0.69	8.80	4.44	15.05	45.31^*	8.01	18.38
LN	0.82	26.66^*	36.11^**	4.94	5.73	5.14	21.42
LP	0.62	35.96^*	40.18^*	6.82	9.16	3.24	4.64
L(C:N)	0.74	28.88^*	33.41^**	4.51	13.25	6.52	13.43
L(C:P)	0.59	34.79^*	33.99^*	8.25	16.55	4.31	2.11
L(N:P)	0.46	22.12	16.76	23.11	9.23	3.60	25.17

叶片化学计量 Leaf stoichiometry	R²	物种多样性解释度 Interpretation of species diversity (%)			土壤化学性质解释度 Interpretation of soil chemical properties (%)
叶片化学计量 Leaf stoichiometry	R²	H′	S	J	SC	SN	SP
LC	0.69	8.80	4.44	15.05	45.31^*	8.01	18.38
LN	0.82	26.66^*	36.11^**	4.94	5.73	5.14	21.42
LP	0.62	35.96^*	40.18^*	6.82	9.16	3.24	4.64
L(C:N)	0.74	28.88^*	33.41^**	4.51	13.25	6.52	13.43
L(C:P)	0.59	34.79^*	33.99^*	8.25	16.55	4.31	2.11
L(N:P)	0.46	22.12	16.76	23.11	9.23	3.60	25.17