科尔沁沙地封育恢复过程中植物群落特征变化及影响因素

doi:10.17521/cjpe.2019.0068

植物生态学报 ›› 2019, Vol. 43 ›› Issue (8): 672-684.DOI: 10.17521/cjpe.2019.0068

科尔沁沙地封育恢复过程中植物群落特征变化及影响因素

王明明^1,²,刘新平^1,^3,^*(),何玉惠⁴,张铜会^1,³,魏静⁵,车力木格^1,²,孙姗姗^1,²

¹中国科学院西北生态环境资源研究院奈曼沙漠化研究站, 兰州 730000
²中国科学院大学, 北京 100049
³中国科学院西北生态环境资源研究院乌拉特荒漠草原研究站, 兰州 730000
⁴中国科学院西北生态环境资源研究院皋兰生态与农业综合研究站, 兰州 730000
⁵北京大学深圳研究生院城市人居环境科学与技术重点实验室, 广东深圳 518000

收稿日期:2019-03-26 修回日期:2019-07-23 出版日期:2019-08-20 发布日期:2020-01-03
通讯作者: 刘新平
基金资助:
国家重点研发计划(2017YFC0506706);国家重点研发计划(2016YFC0500907);国家自然科学基金(41801076);内蒙古自治区科技重大专项(Y749BJ1001);中国科学院科技扶贫项目

How enclosure influences restored plant community changes of different initial types in Horqin Sandy Land

WANG Ming-Ming^1,²,LIU Xin-Ping^1,^3,^*(),HE Yu-Hui⁴,ZHANG Tong-Hui^1,³,WEI Jing⁵,Chelmge ^1,²,SUN Shan-Shan^1,²

¹Naiman Desertification Research Station, Northwest Institute of Eco-Environment and Resources, Chinese Academy of Sciences, Lanzhou 730000, China
²University of Chinese Academy of Sciences, Beijing 100049, China
³Urat Desert-grassland Research Station, Northwest Institute of Eco-Environment and Resources, Chinese Academy of Science, Lanzhou 730000, China
⁴Gaolan Station of Agricultural and Ecological Experiment, Northwest Institute of Eco-Environment and Resources, Chinese Academy of Science, Lanzhou 730000, China
⁵Key Laboratory for Urban Habitat Environmental Science and Technology, Peking University Shenzhen Graduate School, Shenzhen, Guangdong 518000, China

Received:2019-03-26 Revised:2019-07-23 Online:2019-08-20 Published:2020-01-03
Contact: LIU Xin-Ping
Supported by:
National Key R&D Program of China(2017YFC0506706);National Key R&D Program of China(2016YFC0500907);National Natural Science Foundation of China(41801076);Nei Mongol Autonomous Region Science and Technology Major Project(Y749BJ1001);Science and Technology Poverty Alleviation Program of Chinese Academy of Sciences

摘要/Abstract

摘要：

封育是退化沙地植被恢复与生态重建的重要措施, 理解长期处于封育状态下不同类型沙地植物群落特征变化及其影响因素有利于沙地植被恢复和生态重建。该文基于对科尔沁沙地长期封育的流动沙丘(2005年封育)、固定沙丘(1985年封育)和沙质草地(1997年封育)连续多年(2005-2017年)的植物群落调查, 结合土壤种子库、土壤养分以及气象数据, 分析了植物群落特征变化及其对环境变化的响应。研究结果表明流动沙丘植被盖度显著增加, 群落生物量和物种多样性年际间波动变化, 但无明显趋势; 固定沙丘植物群落存在逆行演替趋势, 具体表现为群落生物量、灌木和半灌木以及豆科优势度显著下降, 而一年生和多年生杂类草优势度显著增加; 沙质草地群落物种丰富度和多年生禾草优势度存在降低趋势, 并且一年生杂类草优势度明显高于其他功能群, 群落存在退化现象。3类沙地土壤种子密度变化不显著, 而种子丰富度在流动沙丘显著增加, 在固定沙丘和沙质草地有下降趋势, 土壤养分仅有有效氮和有效磷含量增加。回归分析结果表明气温和降水是影响年内生物量积累的主要因素, 但对年际间群落生物量和物种丰富度变化影响不大。除趋势对应分析结果显示土壤种子库与植物群落之间存在很高的相似性, 典型相关分析结果表明沙质草地植物群落与土壤养分紧密相关, 而固定沙丘群落主要与土壤水分紧密相关。综合以上结果可知, 封育33年的固定沙丘群落和封育21年的沙质草地群落都存在退化现象, 而封育11年的流动沙丘群落正在缓慢恢复, 因此封育年限的设定对退化沙地植被恢复至关重要, 封育时间过长不仅不利于植物群落恢复, 反而会使群落发生逆行演替, 建议封育年限的设定应综合考虑植被退化程度、土壤养分状况、土壤种子库基础以及气候条件等因素的影响。

关键词: 沙地, 封育, 植被恢复, 群落特征, 影响因素

Abstract:

Aims Enclosure is one of the important measures for vegetation restoration of degraded sandy land. Understanding the plant community change of different initial types in long-term state of enclosing is vital for us to understand the vegetation restoration process or re-vegetation in sandy land. This paper aims to analyze the changes of plant communities and its comparative responses to long-term enclosure (2005-2017) of mobile dunes (enclosed in 2005), fixed dunes (enclosed in 1985) and sandy grassland (enclosed in 1997), in relation to soil seed bank, soil nutrient and precipitation and air temperature.
Methods The species composition, height, coverage and above-ground biomass were measured by quadrats in every year during 2005-2017. The soil organic carbon, soil nutrient and soil seed bank were measured by soil cores from the above quadrats in 2008 and 2017. Detrended correspondence analysis (DCA) and canonical correspondence analysis (CCA) were used to explore the relationship between plant community species composition and soil seed bank soil nutrients and soil moisture.
Important findings The results showed that the vegetation coverage and community species richness of mobile dune was significantly increased, but the community biomass had no obvious trend. The biomass of fixed dune community, shrubs, semi-shrubs and the perennial legume functional group dominances decreased significantly, but the dominance of the annual and perennial forbs increased significantly. The annual forbs were the dominant functional group, and community species richness and perennial grass dominances decreased significantly. The soil seed bank had no remarkable change in the three enclosed sandy land communities during 2008-2017, while the seed richness significantly increased in mobile dunes, and that of the fixed dunes and grassland presented downward trends. The soil available nitrogen and available phosphorus were increased significantly. Regression analysis showed the annual variation of plant community biomass was significantly affected by air temperature and precipitation, but there was a little effect on the inter-annual variation of plant community biomass and species richness. Detrended correspondence analysis (DCA) displayed that there was a high similarity between soil seed bank and plant community, and typical correlation analysis (CCA) results showed that grassland community species composition was closely related to soil nutrients. However, fixed dune community species composition was mainly related to soil moisture. In summary, the fixed dune plant community enclosed for 33 years and the sandy grassland plant community enclosed for 21 years present degraded trends, while the mobile dune plant community enclosed for 11 years is slowly recovering, thus, long-term enclosure is not always conducive to the restoration of degraded sandy land vegetation. We suggested that the influence of degradation degree of vegetation, soil nutrients and moisture, soil seed bank and precipitation should be synthetically considered when we set the duration of enclosure for restoration.

Key words: sandy land, enclosure, vegetation restoration, community features, influencing factors

王明明,刘新平,何玉惠,张铜会,魏静,车力木格,孙姗姗. 科尔沁沙地封育恢复过程中植物群落特征变化及影响因素. 植物生态学报, 2019, 43(8): 672-684. DOI: 10.17521/cjpe.2019.0068

WANG Ming-Ming,LIU Xin-Ping,HE Yu-Hui,ZHANG Tong-Hui,WEI Jing,Chelmge ,SUN Shan-Shan. How enclosure influences restored plant community changes of different initial types in Horqin Sandy Land. Chinese Journal of Plant Ecology, 2019, 43(8): 672-684. DOI: 10.17521/cjpe.2019.0068

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2019.0068

https://www.plant-ecology.com/CN/Y2019/V43/I8/672

图/表 9

图1 科尔沁沙地地理位置及研究样地空间分布。

Fig. 1 Geographical location of the Horqin Sandy Land (HSL) and spatial distribution of the sample plots.

图2 科尔沁沙地不同类型封育沙地植物群落特征年际变化(平均值±标准偏差)。FD, 固定沙丘; MD, 流动沙丘; SG, 沙质草地。C, Simpson指数; H, Shannon-Wiener指数; k, 线性回归方程斜率; p, 显著性水平(p < 0.05变化趋势明显)。图中仅列出了置信水平较高的k, R2和p值。

Fig. 2 Interannual changes of plant community features in different enclosure types of sandy lands in the Horqin Sandy Land (mean ± SD). FD, fixed dune; MD, mobile dune; SG, sandy grassland. C, Simpson index; H, Shannon-Wiener index; k, regression line slope; p, statistical significance (p < 0.05 indicates the change trend is significant). Only the k, R2 and p values with higher confidence levels are listed in the figure.

表1 2005和2017年科尔沁沙地不同类型封育沙地植物群落物种组成和物种优势度

Table 1 Species composition and dominance of plant community in different enclosure types of sandy lands in 2005 and 2017

物种 Species	生活型 Biotype	2005			2017
物种 Species	生活型 Biotype	MD	FD	SG	MD	FD	SG
沙蓬 Agriophyllum squarrosum	AF	26.91	0	0	9.60	0	0
苦苣菜 Sonchus oleraceus	AF	10.22	0	0	2.10	0	0
旋覆花 Inula japonica	PF	9.26	0	0	8.75	0	0
狗尾草 Setaria viridis	AG	8.65	2.81	1.52	7.45	3.72	6.72
蒺藜 Tribulus terrestris	AF	5.24	1.00	0	15.77	0	0.92
地梢瓜 Cynanchum thesioides	PF	5.48	0.98	0.84	1.47	2.18	1.88
大果虫实 Corispermum macrocarpum	AF	5.32	1.49	0	3.30	2.12	0
差不嘎蒿 Artemisia halodendron	SH	3.57	11.88	0	5.99	3.20	0
雾冰藜 Bassia dasyphylla	AF	2.69	1.43	0	2.20	1.52	0
马唐 Digitaria sanguinalis	AG	2.45	3.43	0.92	5.95	2.80	0
白草 Pennisetum flaccidum	PG	0	1.92	4.27	0	0	10.55
猪毛菜 Salsola collina	AF	0	0.85	0.85	2.52	7.50	2.17
花苜蓿 Medicago ruthenica	PL	0	11.11	4.41	0	10.50	0
糙隐子草 Cleistogenes squarrosa	PG	0	0	3.58	0	0	3.77
大籽蒿 Artemisia sieversiana	AF	0	0	2.92	0	0	0
地锦 Euphorbia humifusa	AF	0	13.27	0	0.94	1.99	0.85
二裂委陵菜 Potentilla bifurca	AF	0	0	0.71	0	0	0
九顶草 Enneapogon desvauxii	AG	0	0	0	0	0	1.90
兴安乌胡枝子 Lespedeza davurica	SH/PL	0	1.39	2.74	0	2.24	2.83
虎尾草 Chloris virgata	AG	0	0	1.19	0	0	0
画眉草 Eragrostis pilosa	AG	0	3.41	0	1.72	9.99	0
黄蒿 Artemisia scoparia	AF	0	0	22.09	0	6.53	6.11
灰绿藜 Chenopodium glaucum	AF	0	2.46	0.69	0	0	0
鸡眼草 Kummerowia striata	AL	0	0	1.54	0	0	0
尖头叶藜 Chenopodium acuminatum	AF	0	0	0	3.54	4.01	4.94
芦苇 Phragmites australis	PG	0	0	9.56	0	0	6.36
少花米口袋 Gueldenstaedtia verna	PL	0	0	1.58	0	0	0
三芒草 Aristida adscensionis	AG	0	3.07	4.37	0	0	0
砂蓝刺头 Echinops gmelinii	AF	0	1.42	0	0	8.62	0
牻牛儿苗 Erodium stephanianum	AF	0	0	1.67	0	0	12.76
中华苦荬菜 Ixeris chinensis	AF	0	2.17	0	0	0	0
小叶锦鸡儿 Caragana microphylla	SH/PL	0	2.46	0	0	0	0
独行菜 Lepidium apetalum	AG	0	0	0	0	0	5.00
总计 Total	-	10	18	18	14	14	13

表1 2005和2017年科尔沁沙地不同类型封育沙地植物群落物种组成和物种优势度

Table 1 Species composition and dominance of plant community in different enclosure types of sandy lands in 2005 and 2017

物种 Species	生活型 Biotype	2005			2017
物种 Species	生活型 Biotype	MD	FD	SG	MD	FD	SG
沙蓬 Agriophyllum squarrosum	AF	26.91	0	0	9.60	0	0
苦苣菜 Sonchus oleraceus	AF	10.22	0	0	2.10	0	0
旋覆花 Inula japonica	PF	9.26	0	0	8.75	0	0
狗尾草 Setaria viridis	AG	8.65	2.81	1.52	7.45	3.72	6.72
蒺藜 Tribulus terrestris	AF	5.24	1.00	0	15.77	0	0.92
地梢瓜 Cynanchum thesioides	PF	5.48	0.98	0.84	1.47	2.18	1.88
大果虫实 Corispermum macrocarpum	AF	5.32	1.49	0	3.30	2.12	0
差不嘎蒿 Artemisia halodendron	SH	3.57	11.88	0	5.99	3.20	0
雾冰藜 Bassia dasyphylla	AF	2.69	1.43	0	2.20	1.52	0
马唐 Digitaria sanguinalis	AG	2.45	3.43	0.92	5.95	2.80	0
白草 Pennisetum flaccidum	PG	0	1.92	4.27	0	0	10.55
猪毛菜 Salsola collina	AF	0	0.85	0.85	2.52	7.50	2.17
花苜蓿 Medicago ruthenica	PL	0	11.11	4.41	0	10.50	0
糙隐子草 Cleistogenes squarrosa	PG	0	0	3.58	0	0	3.77
大籽蒿 Artemisia sieversiana	AF	0	0	2.92	0	0	0
地锦 Euphorbia humifusa	AF	0	13.27	0	0.94	1.99	0.85
二裂委陵菜 Potentilla bifurca	AF	0	0	0.71	0	0	0
九顶草 Enneapogon desvauxii	AG	0	0	0	0	0	1.90
兴安乌胡枝子 Lespedeza davurica	SH/PL	0	1.39	2.74	0	2.24	2.83
虎尾草 Chloris virgata	AG	0	0	1.19	0	0	0
画眉草 Eragrostis pilosa	AG	0	3.41	0	1.72	9.99	0
黄蒿 Artemisia scoparia	AF	0	0	22.09	0	6.53	6.11
灰绿藜 Chenopodium glaucum	AF	0	2.46	0.69	0	0	0
鸡眼草 Kummerowia striata	AL	0	0	1.54	0	0	0
尖头叶藜 Chenopodium acuminatum	AF	0	0	0	3.54	4.01	4.94
芦苇 Phragmites australis	PG	0	0	9.56	0	0	6.36
少花米口袋 Gueldenstaedtia verna	PL	0	0	1.58	0	0	0
三芒草 Aristida adscensionis	AG	0	3.07	4.37	0	0	0
砂蓝刺头 Echinops gmelinii	AF	0	1.42	0	0	8.62	0
牻牛儿苗 Erodium stephanianum	AF	0	0	1.67	0	0	12.76
中华苦荬菜 Ixeris chinensis	AF	0	2.17	0	0	0	0
小叶锦鸡儿 Caragana microphylla	SH/PL	0	2.46	0	0	0	0
独行菜 Lepidium apetalum	AG	0	0	0	0	0	5.00
总计 Total	-	10	18	18	14	14	13

图3 科尔沁沙地不同类型封育沙地植物群落功能群重要值年际变化。FD, 固定沙丘; MD, 流动沙丘; SG, 沙质草地。AF, 一年生杂类草; AG, 一年生禾草; PF, 多年生杂类草; PG, 多年生禾草; PH, 多年生草本植物, PL, 多年生豆科植物; SH, 灌木或半灌木。k, 回归线斜率。图中仅列出了达到显著性水平(p < 0.05)的检验参数值。

Fig. 3 Interannual variation of the important value of plant community functional groups in different enclosure types of sandy lands in Horqin Sandy Land. FD, fixed dune; MD, mobile dune; SG, sandy grassland. AF, annual forbs; AG; annual grasses; PF, perennial forbs; PG, perennial grasses; PH, perennial grass layer; PL, perennial legumes; SH, shrubs or subshrubs. k, regression line slope. Only the test parameter values that reach the level of significance (p < 0.05) are listed in this figure.

表2 科尔沁沙地不同类型封育沙地植物群落功能群重要值之间的Pearson相关系数

Table 2 Pearson’s correlation coefficient between important values of community functional groups in different types enclosed sandy land in Horqin Sandy Land

沙地 Sandy land	功能群 Functional group	R	p	沙地 Sandy land	功能群 Functional group	R	p
MD	AF vs AG	-0.28	0.35	FD	PL vs SH	0.33	0.28
MD	AF vs SH	-0.21	0.49	SG	AF vs AG	-0.72	<0.01
MD	AG vs SH	-0.11	0.72	SG	AF vs PF	-0.23	0.44
FD	AF vs AG	-0.28	0.36	SG	AF vs PG	-0.81	<0.01
FD	AF vs PF	-0.13	0.67	SG	AF vs PL	-0.17	0.58
FD	AF vs PL	-0.65	<0.05	SG	AG vs PF	0.37	0.21
FD	AF vs SH	-0.71	<0.01	SG	AG vs PG	0.22	0.47
FD	AG vs PF	-0.20	0.51	SG	AG vs PL	-0.14	0.64
FD	AG vs PL	-0.03	0.93	SG	PF vs PG	0.14	0.64
FD	AG vs SH	-0.17	0.57	SG	PF vs PL	-0.04	0.90
FD	PF vs PL	-0.14	0.65	SG	PG vs PL	0.53	0.14
FD	PF vs SH	-0.18	0.57	SG

表3 科尔沁沙地不同类型封育沙地2008和2017年土壤养分、水分、有机质含量以及种子密度和丰富度比较(平均值±标准偏差)

Table 3 Multiple comparisons of soil nutrients, soil moisture, soil organic matter and soil seed density, seed richness in different enclosure types of sandy lands in the Horqin Sandy Land in 2008 and 2017 (mean ± SD)

土壤因子 Soil factor	MD		FD		SG
土壤因子 Soil factor	2008年	2017年	2008年	2017年	2008年	2017年
SOM (g·kg^-1)	0.53 ± 0.08^a	0.42 ± 0.05^b	1.13 ± 0.39^a	1.05 ± 0.36^a	5.64 ± 2.18^a	4.05 ± 0.51^a
TN (g·kg^-1)	0.03 ± 0.05^a	0.024 ± 0.01^a	0.068 ± 0.02^a	0.072 ± 0.06^a	0.34 ± 0.11^a	0.35 ± 0.07^a
AN (mg·kg^-1)	5.35 ± 1.25^b	8.80 ± 2.52^a	6.35 ± 3.20^b	11.28 + 3.60^a	22.51 ± 2.98^b	29.87 ± 4.89^a
AP (mg·kg^-1)	4.14 ± 0.65^b	9.88 ± 0.17^a	3.88 ± 0.92^b	9.87 ± 0.07^a	3.22 ± 1.32^b	9.43 ± 0.11^a
AK (mg·kg^-1)	54.67 ± 7.08^a	43.05 ± 4.60^b	64.58 ± 9.80^a	65.61 ± 11.30^a	116.42 ± 23.30^a	112.47 ± 20.20^a
SSD (粒·m^-2)	492.0 ± 204.40^a	670.0 ± 268.6^a	2 669 ± 1 108^a	1 860 ± 687^a	3 344 ± 1 378^a	4 072 ± 1 639^a
SSR	4.0 ± 1.20^b	5 ± 1.30^a	10 ± 0.50^a	8 ± 1.33^b	12 ± 2.10^a	9 ± 1.24^b
SM (%)	2.43 ± 0.32^b		2.00 ± 0.07^c		3.72 ± 0.10^a

表4 科尔沁沙地不同类型封育沙地植物群落生物量和物种丰富度与气候因子之间关系

Table 4 Relationships between plant community biomass and species richness with climatic factors in different habitats in different types enclosed sandy land in Horqin Sandy Land

气候因子 Climate factor	沙地 Sandy land	生物量 Biomass (g·m^-2)			物种丰富度 Spices richness
气候因子 Climate factor	沙地 Sandy land	方程 Equation	R²	p	方程 Equation	R²	p
年降水量 Annual precipitation (mm)	MD	y = -0.1x + 17.08	0.3	0.98	y = -0.35lg x + 13.34	0.03	0.26
	FD	y = 0.09x + 55.96	0.22	0.45	y = 14.78lg x - 19.67	0.15	0.10
	SG	y = 0.21x + 75.11	0.45	0.13	y = 0.19lg x + 13.50	0.08	0.79
生长季降水量 Growing season precipitation (mm)	MD	y = 0.03x + 8.39	0.13	0.66	y = -0.35lg x + 13.35	0.03	0.27
	FD	y = 0.09x + 58.26	0.21	0.48	y = 14.53lg x - 18.32	0.15	0.11
	SG	y = 0.17x + 93.17	0.32	0.28	y = 0.19lg x + 13.5	0.08	0.79
年平均气温 Annual average air temperature (℃)	MD	y = 1.2x² - 22.64x + 117.04	0.24	0.23	y = -0.35x + 13.34	0.03	0.26
	FD	y = 2.42x² - 39.89x + 234.26	0.42	0.06	y = -0.09x + 16.00	0.09	0.84
	SG	y = 2.65x² - 44.46x + 306.24	0.19	0.35	y = 0.19x + 1.5	0.08	0.79

图4 科尔沁沙地不同类型封育沙地植物群落生物量与生长季降水累积量、积温之间的关系。FD, 固定沙丘; MD, 流动沙丘; SG, 沙质草地。

Fig. 4 Relationships among plant community biomass, and growing season precipitation, growing season accumulated temperature in different enclosure types of sandy land in the Horqin Sandy Land. FD, fixed dune; MD, mobile dune; SG, sandy grassland.

图5 科尔沁沙地不同类型封育沙地植物群落物种、土壤种子库、土壤养分之间的除趋势对应分析(DCA)和典型相关分析(CCA)。A, 群落物种和土壤种子库的DCA分析。B, 群落物种和土壤因子的CCA分析。C, 土壤种子库物种和土壤因子的CCA分析。FD, 固定沙丘; MD, 流动沙丘; SG, 沙质草地。AK, 土壤有效钾; AN, 土壤有效氮; AP, 土壤有效磷; SM, 土壤水分; SOM, 土壤有机质; TN, 土壤全氮。

Fig. 5 Detrended correspondence analysis (DCA) and canonical correspondence analysis (CCA) among plant community species, soil seed bank and soil nutrients. A, the DCA analysis of community species and soil seed bank. B, the CCA analysis of community species and soil nutrients. C, the CCA analysis of soil seed bank species and soil nutrients. FD, fixed dune; MD, mobile dune; SG, sandy grassland. AK, available potassium; AN, available nitrogen; AP, available phosphorus; SM, soil moisture; SOM, soil organic matter; TN, total nitrogen. Veg, vegtation.

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科尔沁沙地封育恢复过程中植物群落特征变化及影响因素

How enclosure influences restored plant community changes of different initial types in Horqin Sandy Land

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