宁夏六盘山华北落叶松人工林植被碳密度时空特征及其环境响应

doi:10.11707/j.1001-7488.LYKX20230037

摘要/Abstract

摘要：

目的: 调查分析宁夏南部六盘山区华北落叶松人工林乔木层、灌木层和草本层碳密度大小及其时空变化规律，建立考虑立地环境和林分结构等主要因子影响的碳密度模型，为我国西北山区复杂地形条件下人工林碳密度估算和可持续经营提供理论基础和决策工具。方法: 采用样地清查与数学模型相结合的方法确定研究区华北落叶松人工林乔木层和林下灌草层碳密度；应用外包线法确定各植被层碳密度对各主要因子（林龄、林分密度、年均降水量、年均气温和坡向）的响应规律及其函数形式，通过连乘耦合构建反映多因子综合影响的植被碳密度模型，并基于实测数据进行模型率定和检验。结果: 1）六盘山区华北落叶松人工林植被碳密度平均为46.82 t·hm^?2，其中乔木层占94.92%，灌木层和草本层分别占4.76%和0.32%。2）乔木层碳密度随林龄、林分密度、年均降水量增大先快速增加、超过阈值后缓慢增加并渐趋最大值，相应阈值为林龄30年、林分密度1 500 株·hm^?2、年均降水量525 mm；随年均气温增加先增后减，6 ℃时达到峰值；随坡向增大（正北为0o，随向两侧偏离正北方向的角度而增大）呈近线性降低。3）林下灌草层碳密度随乔木层碳密度增加近线性减少，随年均气温、年均降水量和坡向的变化规律与乔木层相同，相应阈值同样为年均气温6 ℃、年均降水量525 mm。4）多因子耦合模型可较好估算乔木层和林下灌草层碳密度变化，其R²分别为0.8和0.7。5）根据模型模拟结果，华北落叶松人工林植被碳密度随海拔升高先增后降，垂直差异较大，最高分布范围在海拔2 200~2 400 m之间；受地形和气候影响表现出一定的水平差异，西高东低、南高北低；阴坡植被碳密度高于阳坡，其中乔木层相差约20%，灌草层相差可达1倍。结论: 宁夏六盘山华北落叶松人工林植被碳密度平均为46.82 t·hm^?2，以乔木层为主，受立地环境（年均降水量、年均气温、坡向）和林分结构（林龄、林分密度）的多因子综合影响，林下灌草层碳密度受乔木层生物量影响，应用多因子耦合模型可较好解释并预测其时空变化；合理选择造林立地类型和林分密度有利于维持较好的华北落叶松林植被碳密度和层次组成。

关键词: 华北落叶松人工林, 植被层, 碳密度, 耦合模型, 宁夏六盘山

Abstract:

Objective: In this study, the spatiotemporal variation of vegetation carbon densities (VCD) of the tree layer, understory shrub layer and herb layer of larch (Larix principis-rupprechtii) plantations in the Liupan Mountains area of southern Ningxia of northwest China were investigated and analyzed, and the VCD models considering the effects of multiple factors of site quality and stand structure were established, so as to provide theoretical basis and decision-making tools for estimating VCD and their sustainable management of the larch plantations under complex topographic conditions in the mountainous regions of northwest China. Method: The VCD of tree, shrub and herb layers of larch plantations in the study area was determined by the combined application of sample plot inventory and mathematical models. The upper boundary line method was used to determine the response pattern and corresponding function form of VCD to each main influencing factor (forest age, density, mean annual precipitation, mean annual air temperature, slope aspect). A VCD model that can reflect the comprehensive impact of multiple factors was constructed through multiplicative coupling, and then the measured data were used to calibrate and validate the model. Result: 1) The average VCD of the existing larch plantations in the Liupan Mountains area was 46.82 t C·hm^-2, of which the tree layer accounted for 94.92%, while the shrub layer and herb layer only accounted for 4.76% and 0.32%, respectively. 2) The VCD of tree layer firstly increased rapidly with the increase of forest age, stand density, and mean annual precipitation, and then increased slowly and gradually tended to its maximum value after exceeding the certain threshold values of those factors. The corresponding threshold values for those influencing factors were 30 years of forest age, 1500 tree·hm^-2 of tree density, and 525 mm of mean annual precipitation. The VCD of tree layer firstly increased and then decreased with rising mean annual temperature and reached its peak at 6°C. It decreased nearly linearly with rising slope aspect (0° to due north, increasing with the deviation angle from north direction to both sides). 3) The VCD of the understory shrub-herb layers decreased linearly with rising tree layer VCD, and varied with mean annual temperature and precipitation as well as slope aspect in the same patterns of tree layer VCD and the same threshold values of 6 °C and 525 mm. 4) The established multi-factor coupled model of VCD was able to better estimate the variation of VCD of tree layer and understory shrub-herb layer, with R² of 0.8** and 0.7**, respectively. (5) According to simulation results of the model, the VCD of larch plantations firstly increased and then decreased with rising elevation, showing a relatively bigger difference among elevation gradients, the elevation range with the highest VCD was 2200—2400 m. Due to the effects of topography and climatic factors, there existed a certain horizontal difference, namely decreasing from west to east and from south to north. The VCD on shady slopes was higher than that on sunny slopes, with a difference of about 20% for tree layer and up to 100% for shrub-herb layers. Conclusion: The average VCD of the larch plantations in Liupan Mountains area is 46.82 t C·hm^-2 and mainly composed of tree layer. The VCD is jointly affected by the multiple factors of site quality (precipitation, temperature, slope aspect) and stand structure (age, density). The VCD of understory shrub-herb layers is also influenced by the tree layer biomass. The spatiotemporal variation of VCD of larch plantations can be better explained and predicted by the multi-factor coupled model. The reasonable selection of site types for afforestation and tree density for forest management is beneficial to maintain a better VCD with expected vegetation layer composition for the larch plantations.

Key words: Larix principis-rupprechtii plantations, vegetation layers, carbon density, coupled model, Ningxia Liupan Mountainous

中图分类号:

张紫优,王彦辉,田奥,刘泽彬,郭建斌,于澎涛,王晓,于艺鹏. 宁夏六盘山华北落叶松人工林植被碳密度时空特征及其环境响应[J]. 林业科学, 2023, 59(4): 32-45.

Ziyou Zhang,Yanhui Wang,Ao Tian,Zebin Liu,Jianbin Guo,Pengtao Yu,Xiao Wang,Yipeng Yu. Spatiotemporal Characteristics and Environmental Response of Vegetation Carbon Densities of Larix principis-rupprechtii Plantations in the Liupan Mountains of Ningxia, China[J]. Scientia Silvae Sinicae, 2023, 59(4): 32-45.

图/表 8

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表1

图2

图3

图4

表2

图5

表3

六盘山区不同立地类型华北落叶松人工林植被碳密度变化①"

海拔区间 Elevation ranges /m	坡向 Slope aspect	植被层 Vegetation layer	林分密度Stand density/(tree·hm^?2)														最大值 Max.	最小值 Min.	平均值 Mean
			500					1 000				2 000			3 000
			林龄 Age /a					林龄 Age /a				林龄 Age /a			林龄 Age /a
			10	20	30	40	50	10	20	30	40	10	20	30	10	20
1 800~ 2 000	阴坡 Shady	乔木层 Tree layer	9.5	13.1	20.7	36.9	71.3	16.6	22.9	36.4	64.9	19.2	26.5	42.0	19.4	26.8	71.3	9.5	30.4
		灌草层 Shrub-herb layer	7.4	7.3	7.2	7.0	6.5	6.7	6.6	6.5	6.1	5.5	5.5	5.3	5.3	5.3	7.4	5.3	6.3
		合计 Total	16.8	20.4	27.9	43.9	77.8	23.3	29.6	42.8	71.0	24.8	32.0	47.3	24.8	32.1	77.8	16.8	36.8
	半阴坡 Semi-shady	乔木层 Tree layer	7.8	10.8	17.1	30.5	59.0	13.7	19.0	30.1	53.6	15.9	21.9	34.8	16.1	22.2	59.0	7.8	25.2
		灌草层 Shrub-herb layer	6.1	6.0	6.0	5.8	5.5	5.5	5.5	5.4	5.1	4.6	4.5	4.4	4.4	4.4	6.1	4.4	5.2
		合计 Total	13.9	16.8	23.1	36.3	64.5	19.3	24.5	35.4	58.7	20.5	26.5	39.2	20.5	26.5	64.5	13.9	30.4
	半阳坡 Semi-sunny	乔木层 Tree layer	7.8	10.8	17.1	30.4	58.8	13.7	18.9	30.0	53.5	15.8	21.9	34.7	16.0	22.1	58.8	7.8	25.1
		灌草层 Shrub-herb layer	4.1	4.0	4.0	3.9	3.7	3.7	3.7	3.6	3.4	3.1	3.0	3.0	3.0	2.9	4.1	2.9	3.5
		合计 Total	11.9	14.8	21.1	34.3	62.5	17.4	22.6	33.6	56.9	18.9	24.9	37.6	19.0	25.0	62.5	11.9	28.6
	阳坡 Sunny	乔木层 Tree layer	7.8	10.8	17.1	30.4	58.8	13.7	18.9	30.0	53.5	15.8	21.9	34.7	16.0	22.1	58.8	7.8	25.1
		灌草层 Shrub-herb layer	3.3	3.3	3.3	3.2	3.0	3.0	3.0	2.9	2.8	2.5	2.5	2.4	2.4	2.4	3.3	2.4	2.9
		合计 Total	11.1	14.1	20.3	33.6	61.8	16.7	21.9	32.9	56.3	18.3	24.3	37.1	18.4	24.5	61.8	11.1	28.0
2 000~ 2 200	阴坡 Shady	乔木层 Tree layer	12.9	17.9	28.3	50.5	97.6	22.8	31.4	49.8	88.8	26.3	36.3	57.5	26.6	36.7	97.6	12.9	41.7
		灌草层 Shrub-herb layer	9.6	9.5	9.3	8.9	8.1	8.7	8.5	8.2	7.6	7.2	7.0	6.7	6.9	6.7	9.6	6.7	8.1
		合计 Total	22.5	27.4	37.6	59.4	105.7	31.4	39.9	58.0	96.3	33.5	43.3	64.3	33.5	43.4	105.7	22.5	49.7
	半阴坡 Semi-shady	乔木层 Tree layer	10.7	14.8	23.4	41.7	80.7	18.8	26.0	41.1	73.4	21.7	30.0	47.6	22.0	30.3	80.7	10.7	34.4
		灌草层 Shrub-herb layer	7.9	7.8	7.7	7.5	6.9	7.2	7.1	6.9	6.4	5.9	5.8	5.6	5.7	5.6	7.9	5.6	6.7
		合计 Total	18.6	22.6	31.1	49.2	87.6	26.0	33.0	48.0	79.8	27.7	35.9	53.2	27.7	35.9	87.6	18.6	41.2
	半阳坡 Semi-sunny	乔木层 Tree layer	10.7	14.7	23.3	41.6	80.5	18.8	25.9	41.0	73.2	21.7	29.9	47.4	21.9	30.3	80.5	10.7	34.4
		灌草层 Shrub-herb layer	5.3	5.3	5.2	5.0	4.6	4.8	4.7	4.6	4.3	4.0	3.9	3.8	3.8	3.8	5.3	3.8	4.5
		合计 Total	16.0	20.0	28.5	46.6	85.1	23.6	30.6	45.6	77.5	25.7	33.8	51.2	25.7	34.0	85.1	16.0	38.9
	阳坡 Sunny	乔木层 Tree layer	10.7	14.7	23.3	41.6	80.5	18.8	25.9	41.0	73.2	21.7	29.9	47.4	21.9	30.3	80.5	10.7	34.4
		灌草层 Shrub-herb layer	4.3	4.3	4.2	4.1	3.7	3.9	3.9	3.7	3.5	3.2	3.2	3.1	3.1	3.1	4.3	3.1	3.7
		合计 Total	15.0	19.0	27.6	45.7	84.2	22.7	29.8	44.8	76.7	24.9	33.1	50.5	25.0	33.3	84.2	15.0	38.0
2 200~ 2 400	阴坡 Shady	乔木层 Tree layer	14.6	20.1	31.9	56.8	109.8	25.6	35.3	56.0	99.9	29.6	40.9	64.8	29.9	41.3	109.8	14.6	46.9
		灌草层 Shrub-herb layer	10.4	10.3	10.0	9.6	8.5	9.4	9.2	8.8	8.0	7.7	7.6	7.2	7.4	7.3	10.4	7.2	8.7
		合计 Total	24.9	30.4	41.9	66.4	118.4	35.0	44.5	64.8	107.9	37.3	48.4	71.9	37.3	48.6	118.4	24.9	55.6
	半阴坡 Semi-shady	乔木层 Tree layer	12.0	16.6	26.3	47.0	90.8	21.2	29.2	46.3	82.6	24.5	33.8	53.5	24.7	34.1	90.8	12.0	38.8
		灌草层 Shrub-herb layer	8.6	8.5	8.3	8.0	7.3	7.8	7.6	7.4	6.9	6.4	6.3	6.0	6.2	6.0	8.6	6.0	7.2
		合计 Total	20.6	25.1	34.7	55.0	98.1	28.9	36.9	53.7	89.4	30.9	40.1	59.6	30.9	40.2	98.1	20.6	46.0
	半阳坡 Semi-sunny	乔木层 Tree layer	12.0	16.6	26.3	46.9	90.6	21.1	29.1	46.2	82.4	24.4	33.7	53.4	24.7	34.1	90.6	12.0	38.7
		灌草层 Shrub-herb layer	5.7	5.7	5.6	5.4	4.9	5.2	5.1	4.9	4.6	4.3	4.2	4.0	4.1	4.0	5.7	4.0	4.8
		合计 Total	17.7	22.3	31.9	52.2	95.5	26.3	34.3	51.1	87.0	28.7	37.9	57.4	28.8	38.1	95.5	17.7	43.5
	阳坡 Sunny	乔木层 Tree layer	12.0	16.6	26.3	46.9	90.6	21.1	29.1	46.2	82.4	24.4	33.7	53.4	24.7	34.1	90.6	12.0	38.7
		灌草层 Shrub-herb layer	4.7	4.6	4.5	4.4	4.0	4.2	4.2	4.0	3.7	3.5	3.4	3.3	3.4	3.3	4.7	3.3	3.9
		合计 Total	16.7	21.2	30.8	51.2	94.6	25.3	33.3	50.2	86.1	27.9	37.1	56.7	28.0	37.3	94.6	16.7	42.6
2 400~ 2 600	阴坡 Shady	乔木层 Tree layer	13.8	19.0	30.2	53.8	104.1	24.3	33.5	53.1	94.6	28.0	38.7	61.3	28.4	39.1	104.1	13.8	44.4
		灌草层 Shrub-herb layer	9.1	9.0	8.8	8.4	7.5	8.2	8.1	7.7	7.1	6.8	6.6	6.3	6.5	6.4	9.1	6.3	7.6
		合计 Total	22.9	28.0	39.0	62.2	111.6	32.5	41.5	60.8	101.7	34.8	45.3	67.7	34.8	45.5	111.6	22.9	52.0
	半阴坡 Semi-shady	乔木层 Tree layer	11.4	15.8	25.0	44.5	86.0	20.1	27.7	43.9	78.3	23.2	32.0	50.7	23.4	32.3	86.0	11.4	36.7
		灌木层 Shrub-herb layer	7.5	7.4	7.3	7.0	6.4	6.8	6.7	6.5	6.0	5.6	5.5	5.3	5.4	5.3	7.5	5.3	6.3
		合计 Total	18.9	23.2	32.3	51.5	92.5	26.8	34.4	50.4	84.3	28.8	37.5	56.0	28.8	37.6	92.5	18.9	43.1
	半阳坡 Semi-sunny	乔木层 Tree layer	11.4	15.7	24.9	44.4	85.8	20.0	27.6	43.8	78.1	23.1	31.9	50.6	23.4	32.3	85.8	11.4	36.6
		灌草层 Shrub-herb layer	5.0	5.0	4.9	4.7	4.3	4.5	4.5	4.3	4.0	3.7	3.7	3.6	3.6	3.5	5.0	3.5	4.2
		合计 Total	16.4	20.7	29.8	49.1	90.1	24.6	32.1	48.1	82.1	26.9	35.6	54.1	27.0	35.8	90.1	16.4	40.9
	阳坡 Sunny	乔木层 Tree layer	11.4	15.7	24.9	44.4	85.8	20.0	27.6	43.8	78.1	23.1	31.9	50.6	23.4	32.3	85.8	11.4	36.6
		灌草层 Shrub-herb layer	4.1	4.0	4.0	3.8	3.5	3.7	3.6	3.5	3.3	3.1	3.0	2.9	2.9	2.9	4.1	2.9	3.5
		合计 Total	15.5	19.8	28.9	48.2	89.3	23.7	31.3	47.3	81.3	26.2	34.9	53.5	26.3	35.1	89.3	15.5	40.1
2 600~ 2 800	阴坡 Shady	乔木层 Tree layer	10.3	14.3	22.6	40.4	78.0	18.2	25.1	39.8	71.0	21.0	29.0	46.0	21.3	29.3	78.0	10.3	33.3
		灌草层 Shrub-herb layer	4.9	4.9	4.8	4.7	4.3	4.5	4.4	4.3	4.0	3.7	3.6	3.5	3.6	3.5	4.9	3.5	4.2
		合计 Total	15.3	19.2	27.5	45.0	82.3	22.7	29.5	44.1	75.0	24.7	32.7	49.5	24.8	32.8	82.3	15.3	37.5
	半阴坡 Semi-shady	乔木层 Tree layer	8.6	11.8	18.7	33.4	64.5	15.0	20.8	32.9	58.7	17.4	24.0	38.0	17.6	24.3	64.5	8.6	27.6
		灌草层 Shrub-herb layer	4.1	4.0	4.0	3.9	3.6	3.7	3.7	3.6	3.4	3.1	3.0	2.9	2.9	2.9	4.1	2.9	3.5
		合计 Total	12.6	15.9	22.7	37.3	68.2	18.7	24.4	36.5	62.1	20.5	27.0	41.0	20.5	27.2	68.2	12.6	31.0
	半阳坡 Semi-sunny	乔木层 Tree layer	8.5	11.8	18.7	33.3	64.3	15.0	20.7	32.8	58.5	17.3	23.9	37.9	17.5	24.2	64.3	8.5	27.5
		灌草层 Shrub-herb layer	2.7	2.7	2.6	2.6	2.4	2.5	2.5	2.4	2.3	2.0	2.0	2.0	2.0	1.9	2.7	1.9	2.3
		合计 Total	11.3	14.5	21.3	35.9	66.8	17.5	23.2	35.2	60.8	19.4	26.0	39.9	19.5	26.1	66.8	11.3	29.8
	阳坡 Sunny	乔木层 Tree layer	8.5	11.8	18.7	33.3	64.3	15.0	20.7	32.8	58.5	17.3	23.9	37.9	17.5	24.2	64.3	8.5	27.5
		灌木层 Shrub-herb layer	2.2	2.2	2.2	2.1	2.0	2.0	2.0	1.9	1.9	1.7	1.6	1.6	1.6	1.6	2.2	1.6	1.9
		合计 Total	10.7	14.0	20.8	35.4	66.3	17.0	22.7	34.8	60.4	19.0	25.6	39.5	19.1	25.8	66.3	10.7	29.4
最大值 Max.		乔木层 Tree layer	14.6	20.1	31.9	56.8	109.8	25.6	35.3	56.0	99.9	29.6	40.9	64.8	29.9	41.3	109.8	14.6	46.9
		灌草层 Shrub-herb layer	10.4	10.3	10.0	9.6	8.5	9.4	9.2	8.8	8.0	7.7	7.6	7.2	7.4	7.3	10.4	7.2	8.7
		合计 Total	24.9	30.4	41.9	66.4	118.4	35.0	44.5	64.8	107.9	37.3	48.4	71.9	37.3	48.6	118.4	24.9	55.6
最小值 Min.		乔木层 Tree layer	7.8	10.8	12.3	30.4	58.8	13.7	18.9	30	53.5	15.8	21.9	34.7	16	22.1	58.8	7.8	24.8
		灌草层 Shrub-herb layer	2.2	2.2	2.2	2.1	2.0	2.0	2.0	1.9	1.9	1.7	1.6	1.6	1.6	1.6	2.2	1.6	1.9
		合计 Total	10.7	14.0	20.3	33.6	61.8	16.7	21.9	32.9	56.3	18.3	24.3	37.1	18.4	24.5	61.8	10.7	27.9
平均值 Mean		乔木层 Tree layer	10.6	14.7	22.1	41.4	80.1	18.7	25.8	40.9	72.9	21.6	29.8	47.2	21.8	30.1	80.1	10.6	34.1
		灌草层 Shrub-herb layer	5.9	5.8	5.7	5.5	5.0	5.3	5.2	5.1	4.7	4.4	4.3	4.1	4.2	4.1	5.9	4.1	5.0
		合计 Total	16.5	20.5	28.9	46.9	85.1	24.0	31.0	45.9	77.6	26.0	34.1	51.4	26.0	34.2	85.1	16.5	39.1

表3

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植被层 Vegetation layer	碳密度 Carbon density/（t·hm^?2）	碳密度百分比 Carbon density ratio（%）
乔木层 Tree layer	44.44±27.63a	94.92±10.65a
灌木层 Shrub layer	2.23±2.13b	4.76±3.46b
草本层 Herb layer	0.15±0.15c	0.32±0.25c
合计 Total	46.82±28.13	100

植被层 Vegetation layer	多因子耦合模型 Multi-factor coupling models	R²		NSE
植被层 Vegetation layer	多因子耦合模型 Multi-factor coupling models	拟合 Calibration	验证 Validation	拟合 Calibration	验证 Validation
乔木层 Tree layer	$\begin{gathered} [15.92 + 3.83 \times \exp ( - 0.075 \times {\rm{age}})] \times [0.44 - 0.80 \times \exp ( - 2.53 \times {10^{ - 3} } \times N)] \times \\ ( - 2.92 \times {\rm{MA}}{{\rm{T}}^2} + 34.62 \times {\rm{MAT}} - 48.09) \times [0.30 - 6.48 \times \exp ( - 7.41 \times {10^{ - 3} } \times {\rm{MAP}})] \times \\ [0.20 + 0.042 \times \exp ( - 0.097 \times {\rm{asp}})] \\ \end{gathered}$	0.80	0.77	0.68	0.68
灌草层 Shrub-herb layers	$\begin{gathered} [ - 0.68 + 1.39 \times \exp ( - 0.65 \times {\rm{TBC}})] \times ( - 0.61 \times {\rm{MA}}{{\rm{T}}^2} + 7.82 \times {\rm{MAT}} - 15.61) \times \\ [ - 0.40 + 0.40 \times \exp (3.04 \times {10^{ - 6} } \times {\rm{MAP}})] \times [ - 0.35 + 6.93 \times \exp ( - 4.13 \times {10^{ - 3} } \times {\rm{asp}})] \\ \end{gathered}$	0.70	0.66	0.66	0.64

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