Effects of exclosure on community inter-specific relationships in a typical temperate grassland

doi:10.17521/cjpe.2015.0446

Abstract

Abstract:

Aims This study aimed to understand the impact of exclosure on interspecific associations of species in a typical temperate grassland. We compared the species associations of dominant and companion species in grassland plots with the six-year exclosure and grazing in Taips County, Nei Mongol.Methods We selected 14 species according to important values in each plot, using variance ratio to analyze the overall association. The χ² test with 2 × 2 contingency table, Ochiai index and spearman rank correlation test were applied to analyze the inter-specific associations of dominant and companion species. We used the niche breadth index to analyze the ability of resource utilization of species and inter-specific competition for grazed plots and ones with exclosure.Important findings The inter-specific associations among 91 species pairs inside the exclosure showed that nine species pairs were significantly positive but 4 pairs were negative based on the χ² test with 2 × 2 contingency table. Ten species pairs had significantly positive correlations and 16 species pairs were negative through the Spearman correlation test. For the plots outside the exclosure, we found 19 species pairs were positively associated but 8 pairs were negative associated using the χ² test, and 12 species pairs were significantly positively correlated but 30 pairs were negatively correlated using the Spearman correlation test. The results indicated that the community inside the exclosure was in a continuous development state while the outside community was degradating. The Ochiai indices of dominant species were high both inside and outside the exclosure but the differences using the χ²test were nonsignificant. It implied that there was some connection among dominant species but their patterns of distribution were relatively independent. The niche breadth for dominant species inside the exclosure was higher than ones outside, which suggested the exclosure has an important role in protecting dominant species.

http://jtp.cnki.net/bilingual/detail/html/ZWSB201606004

Key words: exclosure, importance value, interspecific association, niche breadth, resources utilization

Rong GONG, Qiong GAO, Ya-Lin WANG. Effects of exclosure on community inter-specific relationships in a typical temperate grassland[J]. Chin J Plant Ecol, 2016, 40(6): 554-563.

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URL: https://www.plant-ecology.com/EN/10.17521/cjpe.2015.0446

https://www.plant-ecology.com/EN/Y2016/V40/I6/554

Figures/Tables 10

Table 1 The frequency, importance value and niche breadth of species inside the exclosure

编号 Numbers	物种 Species	频度 Frequency	重要值 Importance value	生态位宽度 Niche breadth
1	猪毛蒿 Artemisia scoparia	98	9.814	1.95
2	冷蒿 Artemisia frigida	84	8.608	1.88
3	羊草 Leymus chinensis	91	7.747	1.93
4	糙隐子草 Cleistogenes squarrosa	81	6.601	1.96
5	克氏针茅 Stipa krylovii	70	5.689	1.82
6	花苜蓿 Medicago ruthenica	53	5.101	1.72
7	碱韭 Allium polyrhizum	67	4.243	1.81
8	二裂委陵菜 Potentilla bifurca	45	3.965	1.47
9	龙蒿 Artemisia dracunculus	23	3.506	1.24
10	大籽蒿 Artemisia sieversiana	8	2.581	0.86
11	寸草 Carex duriuscula	15	2.553	1.14
12	冰草 Agropyron cristatum	7	2.529	0.84
13	阿尔泰狗娃花 Aster altaicus	17	2.385	1.22
14	瓣蕊唐松草 Thalictrum petaloideum	6	2.003	0.75

Table 2 The frequency, importance value and niche breadth of species outside the exclosure

编号 Numbers	物种 Species	频度 Frequency	重要值 Importance value	生态位宽度 Niche breadth
1	糙隐子草 Cleistogenes squarrosa	97	10.180	1.86
2	冷蒿 Artemisia frigida	79	9.674	1.84
3	猪毛蒿 Artemisia scoparia	92	8.676	1.93
4	羊草 Leymus chinensis	73	6.447	1.81
5	克氏针茅 Stipa krylovii	55	6.284	1.71
6	碱韭 Allium polyrhizum	82	5.518	1.89
7	栉叶蒿 Neopallasia pectinata	20	4.491	1.25
8	二裂委陵菜 Potentilla bifurca	33	3.874	1.61
9	大籽蒿 Artemisia sieversiana	20	3.657	1.29
10	冰草 Agropyron cristatum	26	3.397	1.41
11	寸草 Carex duriuscula	28	3.165	1.42
12	花苜蓿 Medicago ruthenica	37	2.864	1.55
13	天名精 Carpesium abrotanoides	22	2.613	1.32
14	龙蒿 Artemisia dracunculus	18	2.585	1.34

Table 3 Dynamics of overall associations inside and outside the exclosure

	方差比率 Variance ratio	检验统计量 Statistic ω	χ²临界值 χ² threshold value	结果 Results
围栏内 Inside the exclosure	1.320	132.42	[77.93, 124.34]	显著正联结 Significantly positive association
围栏外 Outside the exclosure	0.931	93.11	[77.93, 124.34]	不显著负联结 Insignificantly negative association

Fig. 1 The semi-matrix of the χ2 test inside the exclosure. Species codes: 1, Artemisia scoparia; 2, Artemisia frigid; 3, Leymus chinensis; 4, Cleistogenes squarrosa; 5, Stipa krylovii; 6, Medicago ruthenica; 7, Allium polyrhizum; 8, Potentilla bifurca; 9, Artemisia dracunculus; 10, Artemisia sieversiana; 11, Carex duriuscula; 12, Agropyron cristatum; 13, Aster altaicus; 14, Thalictrum petaloideum.

Fig. 2 The semi-matrix of the χ2 test outside the exclosure. Species codes: 1, Artemisia scoparia; 2, Artemisia frigid; 3, Leymus chinensis; 4, Cleistogenes squarrosa; 5, Stipa krylovii; 6, Medicago ruthenica; 7, Allium polyrhizum; 8, Potentilla bifurca; 9, Artemisia dracunculus; 10, Artemisia sieversiana; 11, Carex duriuscula; 12, Agropyron cristatum; 13, Aster altaicus; 14, Thalictrum petaloideum.

Table 4 Ochiai indices (OI) of inter-specific associations inside and outside the exclosure

	0 ≤ OI < 0.2		0.2 ≤ OI < 0.4		0.4 ≤ OI < 0.6		0.6 ≤ OI < 0.8		0.8 ≤ OI < 1
	种对数 Species pairs	百分比 Percentage (%)	种对数 Species pairs	百分比 Percentage (%)	种对数 Species pairs	百分比 Percentage (%)	种对数 Species pairs	百分比 Percentage (%)	种对数 Species pairs	百分比 Percentage (%)
围栏内 Inside the exclosure	20	22	36	39.6	15	16.5	7	7.7	13	14.3
围栏外 Outside the exclosure	8	8.8	21	23.1	46	50.5	8	8.8	9	9.9

Fig. 3 The semi-matrix of Ochiai indices (OI) for species in plots inside the exclosure. Species codes: 1, Artemisia scoparia; 2, Artemisia frigid; 3, Leymus chinensis; 4, Cleistogenes squarrosa; 5, Stipa krylovii; 6, Medicago ruthenica; 7, Allium polyrhizum; 8, Potentilla bifurca; 9, Artemisia dracunculus; 10, Artemisia sieversiana; 11, Carex duriuscula; 12, Agropyron cristatum; 13, Aster altaicus; 14, Thalictrum petaloideum.

Fig. 4 The semi-matrix of Ochiai indices (OI) for species in plots outside the exclosure. Species codes: 1, Artemisia scoparia; 2, Artemisia frigid; 3, Leymus chinensis; 4, Cleistogenes squarrosa; 5, Stipa krylovii; 6, Medicago ruthenica; 7, Allium polyrhizum; 8, Potentilla bifurca; 9, Artemisia dracunculus; 10, Artemisia sieversiana; 11, Carex duriuscula; 12, Agropyron cristatum; 13, Aster altaicus; 14, Thalictrum petaloideum.

Fig. 5 The semi-matrix of the Spearman correlation analysis for species in plots inside the exclosure. Species codes: 1, Artemisia scoparia; 2, Artemisia frigid; 3, Leymus chinensis; 4, Cleistogenes squarrosa; 5, Stipa krylovii; 6, Medicago ruthenica; 7, Allium polyrhizum; 8, Potentilla bifurca; 9, Artemisia dracunculus; 10, Artemisia sieversiana; 11, Carex duriuscula; 12, Agropyron cristatum; 13, Aster altaicus; 14, Thalictrum petaloideum.

Fig. 6 The semi-matrix of the Spearman correlation analysis for species in plots outside the exclosure. Species codes: 1, Artemisia scoparia; 2, Artemisia frigid; 3, Leymus chinensis; 4, Cleistogenes squarrosa; 5, Stipa krylovii; 6, Medicago ruthenica; 7, Allium polyrhizum; 8, Potentilla bifurca; 9, Artemisia dracunculus; 10, Artemisia sieversiana; 11, Carex duriuscula; 12, Agropyron cristatum; 13, Aster altaicus; 14, Thalictrum petaloideum.

References 28

1	Aguiar MR, Sala OE (1994). Competition, facilitation, seed distribution and the origin of patches in a Patagonian steppe.Oikos, 70, 26-34.
2	Callaway RM, Brooker RW, Choler P, Kikvidze Z, Lortie CJ, Michalet R, Paolini L, Pugnaire FI, Newingham B, Aschehoug ET, Armas C, Kikodze D, Cook BJ (2002). Positive interactions among alpine plants increase with stress.Nature, 417, 844-848.
3	Callaway RM (1996). Competition and facilitation: Contrasting effects of Artemisia tridentata on desert vs montane pines.Ecology, 77, 2130-2141.
4	Deng ZX, Liu JY, Zhan JY, Zhao T (2004). Dynamic simulation on the spatio-temporal patterns of land use change in Taibus County.Chinese Journal of Plant Ecology, (2), 147-156.(in Chinese with English abstract)[邓祥征, 刘纪远, 战金艳, 赵涛 (2004). 太仆寺旗土地利用变化时空格局的动态模拟. 地理研究, (2), 147-156.]
5	Grechanichenko TE (2001). Changes in the Carabid fauna (Coleoptera, Carabidae) of meadow steppes during the past decades.Russian Journal of Ecology, 32, 117-121.
6	Henry GHR (1998). Environmental influences on the structure of sedge meadows in the canadian high arctic.Plant Ecology, 134, 119-129.
7	Jia CXZ, Li SF, Su JR, Su L (2014). Effects of selective logging on inter- and intra-specific interactions of trees in a natural Pinus kesiya forest.Chinese Journal of Plant Ecology, 38, 1296-1306. (in Chinese with English abstract)[贾呈鑫卓, 李帅锋, 苏建荣, 苏磊 (2014). 择伐对思茅松天然林乔木种间与种内关系的影响. 植物生态学报, 38, 1296-1306.]
8	Jing GH, Cheng JM, Su JS, Wei L, Shi XX, Jin JW (2015). Response of dominant population niche breadths and niche overlaps to various disturbance factors in typical steppe fenced grassland of China’s Loess Plateau region.Acta Prataculturae Sinica, 24(9), 43-52. (in Chinese with English abstract)[井光花, 程积民, 苏纪帅, 魏琳, 史晓晓, 金晶炜 (2015). 黄土区长期封育草地优势物种生态位宽度与生态位重叠对不同干扰的响应特征. 草业学报, 24(9), 43-52.]
9	Li JB, Ma CP, Lu WH, Zhu JZ, Tursunay, Reyimjan (2009). Effects of enclosing spring-autumn grassland on aboveground biomass at Zhaosu horse ranch.Grassland and Turf, (2), 46-50. (in Chinese with English abstract)[李军保, 马存平, 鲁为华, 朱进忠, 吐尔逊娜依, 热依木江 (2009). 围栏封育对昭苏马场春秋草地地上植物量的影响. 草原与草坪, (2), 46-50.]
10	Li LL, Xu N, Mo XQ, Li HY (2014). Numerical analysis of plants inter-specific relationships in Qilihai wetland. Bulletin of Soil and Water Conservation, 34(4), 70-75. (in Chinese with English abstract)[李兰兰, 许诺, 莫训强, 李洪远 (2014). 七里海湿地植物种间关系的数量分析. 水土保持通报, 34(4), 70-75.]
11	Li XR (1999). Inter-specific association and correlation of shrub layer in the coniferous-broad leaves mixed geobotanical zone of Russia Plain.Acta Ecologica Sinica, (1), 57-62. (in Chinese with English abstract)[李新荣 (1999). 俄罗斯平原针阔混交林群落的灌木层植物种间相关研究. 生态学报, (1), 57-62.]
12	Li ZL, Qin WH, ZHou SB, ZHang JB, Wang Z, Wang YJ (2014). Study on plant niche under fencing measures in the semi-arid grassland of North China.Acta Agrectir Sinica, 22, 1186-1193. (in Chinese with English abstract)[李中林, 秦卫华, 周守标, 张纪兵, 汪贞, 王运静 (2014). 围栏封育下华北半干旱草原植物生态位研究. 草地学报, 22, 1186-1193.]
13	Liu JH, Gao H, Zhang LH, Chen LP, Zhao NX, Gao YB (2010). Comparative analysis of inter-specific association within the Stipa grandis-S. krylovii community in typical steppe of Inner Mongolia, China.Chinese Journal of Plant Ecology, 34, 1016-1024. (in Chinese with English abstract)[刘珏宏, 高慧, 张丽红, 陈丽萍, 赵念席, 高玉葆 (2010). 内蒙古锡林郭勒草原大针茅-克氏针茅群落的种间关联特征分析. 植物生态学报, 34, 1016-1024.]
14	Liu JL (2013). Study on Impact of Mowing and Grazing on Vegetation and Soil of Typical Steppe in Xilingol, Inner Mongolia. PhD dissertation, Chinese Academy of Agricultural Sciences, Beijing. 91. (in Chinese with English abstract)[柳剑丽 (2013). 刈割与放牧对锡林郭勒典型草原植被和土壤影响的研究. 博士学位论文, 中国农业科学院, 北京. 91.]
15	Liu XD, Zhang KB, Wang LL, Yang XH (2015). How enclosure affects community characteristics of the sandy grassland in semi-arid areas of northwestern China.Journal of Beijing Forestry University, 37(2), 48-54. (in Chinese with English abstract)[刘小丹, 张克斌, 王黎黎, 杨晓晖 (2015). 封育对半干旱区沙化草地群落特征的影响. 北京林业大学学报, 37(2), 48-54.]
16	Lou YJ, Zhao KY (2008). Analysis inter-specific associations of Carex lasiocarpa community in recent 30-years succession in Sanjiang Plain.Chinese Journal of Ecology, 27, 509-513. (in Chinese with English abstract)[娄彦景, 赵魁义 (2008). 三江平原毛苔草群落近30年演替过程中的种间联结性分析. 生态学杂志, 27, 509-513.]
17	Lü SJ, Liu HM, Wu YL, Wei ZJ, Sun SX, Wang YJ, Lin S (2014). Effect of grazing on niche of constructive and dominant species in the desert steppe.Ecology and Environmental Sciences, 23, 1600-1609. (in Chinese with English abstract)[吕世杰, 刘红梅, 吴艳玲, 卫智军, 孙世贤, 王颖杰, 林森 (2014). 放牧对荒漠草原建群种和优势种生态位的影响. 生态环境学报, 23, 1600-1609.]
18	Nathan R (2006). Long-distance dispersal of plants.Science, 313, 786-788.
19	Nuñez CI, Aizen MA, Ezcurra C (1999). Species associations and nurse plant effects in patches of high-Andean vegetation.Journal of Vegetation Science, 10, 357-364.
20	Palmer TM, Stanton ML, Young TP (2003). Competition and Coexistence: Exploring mechanisms that restrict and maintain diversity within mutualist guilds.Chicago Journals,162, s63-s79.
21	Wang J (2003). Studies on Response of Artemisia frigida Population to Grazing Disturbance. PhD dissertation, Mongolia University, Hohhot. 93. (in Chinese with English abstract)[王静 (2003). 冷蒿种群对放牧干扰响应的研究. 博士学位论文, 内蒙古大学, 呼和浩特. 93.]
22	Wang SP, Wang YF (2001). Study on over-compensation growth of Cleistogenes squarrosa population in Inner Mongolia Steppe.Acta Botanica Sinica, 43, 413-418. (in Chinese with English abstract)[汪诗平, 王艳芬 (2001). 不同放牧率下糙隐子草种群补偿性生长的研究. 植物学报, 43, 413-418.]
23	Xu L, Gao Q, Wang YL (2014). Species richness within a six-year slope exclosure in a temperate grassland and its relationship with aboveground biomass.Ecology and Environmental Sciences, 23, 398-405. (in Chinese with English abstract)[徐粒, 高琼, 王亚林 (2014). 围封6年对温带典型草原坡地物种多样性及其与地上生物量的关系的影响. 生态环境学报, 23, 398-405.]
24	Zaal K, Georgi N (1998). Facilitation in subnival vegetation patches.Journal of Vegetation Science, 9, 261-264.
25	Zhang F, Shangguan TL (2000). Numerical analysis of interspecific relationships in Anelawagnus mollis community in Shanxi.Acta Phytoecologica Sinica, 24, 351-355. (in Chinese with English abstract)[张峰, 上官铁梁 (2000). 山西翅果油树群落种间关系的数量分析. 植物生态学报, 24, 351-355.]
26	Zhang JT (2004). Quantitative Ecology. Science Press, Beijing. (in Chinese)[张金屯 (2004). 数量生态学. 科学出版社, 北京.]
27	Zhang MX, Wang DX, Kang B, Zhang GG, Liu P, Du YL, Yu F (2015). Interspecific associations of dominant plant populations in secondary forest of Pinus armandii in Qinling Mountains.Scientia Silvae Sinicae, 51(1), 12-21. (in Chinese with English abstract)[张明霞, 王得祥, 康冰, 张岗岗, 刘璞, 杜焰玲, 于飞 (2015). 秦岭华山松天然次生林优势种群的种间联结性. 林业科学, 51(1), 12-21.]
28	Zhao DL (2010). Community Ecological Effect of Grazing on Stipa krylovii Steppe. Master degree dissertation, Inner Mongolia University, Hohhot. 54. (in Chinese with English abstract).[赵登亮 (2010). 放牧对克氏针茅草原群落的生态效应. 硕士学位论文, 内蒙古大学, 呼和浩特. 54.]

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