平茬处理对小叶锦鸡儿灌丛邻居植物群落空间格局的影响
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摘要
近几十年来,内蒙古典型草原小叶锦鸡儿(Caragana microphylla)灌丛化现象普遍发生,严重影响了草地生产力。平茬作为灌丛化草地主要管理方式之一,已受到广泛关注;然而平茬如何影响灌丛邻居植物群落的格局动态及其下土壤理化性质变化,仍缺少相应研究。以内蒙古草原小叶锦鸡儿灌丛化草地为研究对象,进行平茬处理,并依据距离小叶锦鸡儿灌丛的远近位置(远—2.5 m,近—0.5 m)设置样方进行植物群落调查和土壤样品的采集,经分析发现:(1)平茬处理显著提高了灌丛邻居植物群落均匀度及邻居植物群落下土壤全碳、全氮含量。(2)借助零模型(Null-model)分析,得出未平茬处理条件下,距离灌丛远近两种位置群落均表现为竞争性格局;而平茬处理条件下,群落竞争性格局作用弱化,且距离灌丛2.5 m位置处群落表现为促进性格局。(3)调查群落中物种间的关系多为中性作用;仅有少数物种对表现为显著正(或负)相互作用关系,且多由群落中优势物种组成;平茬处理条件下群落中显著负相互作用关系物种对比例下降,支持群落整体竞争性格局弱化这一结论。所得结果不仅为小叶锦鸡儿灌丛平茬处理条件下邻居植物群落及土壤环境的动态变化的探讨提供了实验依据,而且对灌丛化草地的恢复具有一定的实践指导意义。
Over recent decades, the Inner Mongolia Steppe has degraded seriously owing to global climate change and extreme human activities, and the population of Caragana microphylla(a native leguminous shrub) spread quickly in the degraded regions. To relieve the pressure of shrub-encroachment and promote the restoration of degraded grasslands, several measures, such as pruning, have been carried out. However, little is known about how pruning affects the community patterns and soil physical and chemical properties. In the present study, we selected a C. microphylla-dominated grassland in Zhenglan Banner, Inner Mongolia Autonomous Region, and set up pruning and non-pruning(control) plots with a combination of sites at different distances to the shrub edge(0.5 and 2.5 m distances, named 0.5 and 2.5 m-site), as treatments to investigated the soil characteristics and spatial distribution patterns of neighbouring plant communities by Null-model. The results were as follows.(1) Soil total carbon, soil total nitrogen, and Pielou evenness index were significantly(P < 0.05) influenced by pruning treatments, but not by sites, being significantly higher under pruning treatments than those under non-pruning treatments;(2) Under non-pruning treatments, the community spatial distribution patterns were dominated by competition; however, under pruning treatments, community spatial distribution patterns showed less competition, and the 2.5 m-site under the pruning treatments was dominated by facilitation;(3) Only a few pairwise species were detected to have significant negative or positive interactions(the Standardized Effect Size was greater than 2 or less than-2), and they consisted of the dominant species of their communities. However, they still played a key role in explaining community spatial distribution patterns. The number of pairwise species that have significant negative interactions under pruning treatments was less than that under non-pruning treatments, suggesting less competition in pruning communities. The results not only provided an experimental basis for exploring variations in community spatial distribution patterns by pruning C. microphylla, but also provided practical guidance for the restoration of shrub-encroached grasslands.
Over recent decades, the Inner Mongolia Steppe has degraded seriously owing to global climate change and extreme human activities, and the population of Caragana microphylla(a native leguminous shrub) spread quickly in the degraded regions. To relieve the pressure of shrub-encroachment and promote the restoration of degraded grasslands, several measures, such as pruning, have been carried out. However, little is known about how pruning affects the community patterns and soil physical and chemical properties. In the present study, we selected a C. microphylla-dominated grassland in Zhenglan Banner, Inner Mongolia Autonomous Region, and set up pruning and non-pruning(control) plots with a combination of sites at different distances to the shrub edge(0.5 and 2.5 m distances, named 0.5 and 2.5 m-site), as treatments to investigated the soil characteristics and spatial distribution patterns of neighbouring plant communities by Null-model. The results were as follows.(1) Soil total carbon, soil total nitrogen, and Pielou evenness index were significantly(P < 0.05) influenced by pruning treatments, but not by sites, being significantly higher under pruning treatments than those under non-pruning treatments;(2) Under non-pruning treatments, the community spatial distribution patterns were dominated by competition; however, under pruning treatments, community spatial distribution patterns showed less competition, and the 2.5 m-site under the pruning treatments was dominated by facilitation;(3) Only a few pairwise species were detected to have significant negative or positive interactions(the Standardized Effect Size was greater than 2 or less than-2), and they consisted of the dominant species of their communities. However, they still played a key role in explaining community spatial distribution patterns. The number of pairwise species that have significant negative interactions under pruning treatments was less than that under non-pruning treatments, suggesting less competition in pruning communities. The results not only provided an experimental basis for exploring variations in community spatial distribution patterns by pruning C. microphylla, but also provided practical guidance for the restoration of shrub-encroached grasslands.
引文
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[4] Tilman D.Resource Competition and Community Structure.Monographs in population biology,1982,17(17):1-296.
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[9] 韩砚君,牛建明,张庆,董建军,张雪峰,康萨如拉.锡林河流域近30年草原植被格局动态及驱动力分析.中国草地学报,2014,36(2):70- 77.
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[12] 蔡文涛,来利明,李贺祎,周继华,管天玉,张晓龙,高楠楠,郑元润.草地灌丛化研究进展.应用与环境生物学报,2016,22(4):531- 537.
[13] 杨永胜,卜崇峰,高国雄.平茬措施对柠条生理特征及土壤水分的影响.生态学报,2012,32(4):1327- 1336.
[14] 张海娜,方向文,蒋志荣,冯彦皓.柠条平茬处理后不同组织游离氨基酸含量.生态学报,2011,31(9):2454- 2460.
[15] 牛钰杰,杨思维,王贵珍,刘丽,花立民.放牧干扰下高寒草甸物种多样性指数评价与选择.应用生态学报,2017,28(6):1824- 1832.
[16] Gotelli N J.Null model analysis of species co-occurrence patterns.Ecology,2000,81(9):2606- 2621.
[17] Zhang J,Hao Z Q,Song B,Li B H,Wang X G,Ye J.Fine-scale species co-occurrence patterns in an old-growth temperate forest.Forest Ecology and Management,2009,257(10):2115- 2120.
[18] Stone L,Roberts A.The checkerboard score and species distributions.Oecologia,1990,85(1):74- 79.
[19] López R P,Valdivia S,Rivera M L,Rios R S.Co-occurrence patterns along a regional aridity gradient of the subtropical Andes do not support stress gradient hypotheses.PLoS One,2013,8(3):e58518.
[20] Gotelli N J,Ulrich W.The empirical bayes approach as a tool to identify non-random species associations.Oecologia,2010,162(2):463- 477.
[21] Veech J A,Miguel Araújo.The pairwise approach to analysing species co-occurrence.Journal of Biogeography,2014,41(6):1029-1035
[22] 陈蕾伊,沈海花,方精云.灌丛化草原:一种新的植被景观.自然杂志,2014,36(6):391- 396.
[23] Xu Z W,Wan S Q,Ren H Y,Han X G,Li M H,Cheng W X,Jiang Y.Effects of water and nitrogen addition on species turnover in temperate grasslands in northern China.PLoS One,2012,7(6):e39762.
[24] 辛小娟.氮、磷添加对亚高山草甸地上/地下生物量分配及植物功能群组成的影响[D].兰州:兰州大学,2011.
[25] 陈慧敏,石福习,杨桂生,张新厚,毛瑢.养分添加对三江平原沼泽化草甸植物群落组成和地上生物量的影响.生态学杂志,2016,35(6):1440- 1446.
[26] Blaser W J,Sitters J,Hart S P,Edwards P J,Olde Venterink H.Facilitative or competitive effects of woody plants on understorey vegetation depend on N-fixation,canopy shape and rainfall.Journal of Ecology,2013,101(6):1598- 1603.
[27] Dohn J,Dembélé F,Karembé M,Moustakas A,Amévor K A,Hanan N P.Tree effects on grass growth in savannas:competition,facilitation and the stress-gradient hypothesis.Journal of Ecology,2013,101(1):202- 209.
[28] Howard K S C,Eldridge D J,Soliveres S.Positive effects of shrubs on plant species diversity do not change along a gradient in grazing pressure in an arid shrubland.Basic and Applied Ecology,2012,13(2):159- 168.
[29] Jeltsch F,Milton S J,Dean W R J,Van Rooyen N,Moloney K A.Modelling the impact of small-scale heterogeneities on tree-grass coexistence in Semi-Arid Savannas.Journal of Ecology,1998,86(5):780- 793.
[30] 彭海英,李小雁,童绍玉.干旱半干旱区草原灌丛化研究进展.草业学报,2014,23(2):313- 322.
[31] 高玉葆,任安芝,王巍,王金龙.科尔沁沙地黄柳再生枝与现存枝形态和光合特征的比较.生态学报,2002,22(10):1758- 1764.
[32] Zhang G F,Zhao W Z.Species-specific traits determine shrub-annual interactions during a growing season.Journal of Arid Land,2015,7(3):403- 413.
[33] Sfenthourakis S,Tzanatos E,Giokas S.Species co-occurrence:the case of congeneric species and a causal approach to patterns of species association.Global Ecology and Biogeography,2006,15(1):39- 49.
[34] Arita H T.Species co-occurrence analysis:pairwise versus matrix-level approaches.Global Ecology and Biogeography,2016,25(11):1397- 1400.
[2] Eldridge D J,Bowker M A,Maestre F T,Roger E,Reynolds J F,Whitford W G.Impacts of shrub encroachment on ecosystem structure and functioning:towards a global synthesis.Ecology Letters,2011,14(7):709- 722.
[3] 张金屯.植物种群空间分布的点格局分析.植物生态学报,1998,22(4):344- 349.
[4] Tilman D.Resource Competition and Community Structure.Monographs in population biology,1982,17(17):1-296.
[5] Bertness M D,Callaway R.Positive interactions in communities.Trends in Ecology & Evolution,1994,9(5):191- 193.
[6] Bruno J F,Stachowicz J J,Bertness M D.Inclusion of facilitation into ecological theory.Trends in Ecology & Evolution,2003,18(3):119- 125.
[7] Gotelli N J.Research frontiers in null model analysis.Global Ecology and Biogeography,2001,10(4):337- 343.
[8] Kang L,Han X G,Zhang Z B,Sun O J.Grassland ecosystems in China:review of current knowledge and research advancement.Philosophical Transactions of the Royal Society B:Biological Sciences,2007,362(1482):997- 1008.
[9] 韩砚君,牛建明,张庆,董建军,张雪峰,康萨如拉.锡林河流域近30年草原植被格局动态及驱动力分析.中国草地学报,2014,36(2):70- 77.
[10] Ratajczak Z,Nippert J B,Collins S L.Woody encroachment decreases diversity across North American grasslands and savannas.Ecology,2012,93(4):697-703.
[11] Eldridge D J,Maestre F T,Maltez-Mouro S,Bowker M A.A global database of shrub encroachment effects on ecosystem structure and functioning.Ecology,2012,93(11):2499- 2499.
[12] 蔡文涛,来利明,李贺祎,周继华,管天玉,张晓龙,高楠楠,郑元润.草地灌丛化研究进展.应用与环境生物学报,2016,22(4):531- 537.
[13] 杨永胜,卜崇峰,高国雄.平茬措施对柠条生理特征及土壤水分的影响.生态学报,2012,32(4):1327- 1336.
[14] 张海娜,方向文,蒋志荣,冯彦皓.柠条平茬处理后不同组织游离氨基酸含量.生态学报,2011,31(9):2454- 2460.
[15] 牛钰杰,杨思维,王贵珍,刘丽,花立民.放牧干扰下高寒草甸物种多样性指数评价与选择.应用生态学报,2017,28(6):1824- 1832.
[16] Gotelli N J.Null model analysis of species co-occurrence patterns.Ecology,2000,81(9):2606- 2621.
[17] Zhang J,Hao Z Q,Song B,Li B H,Wang X G,Ye J.Fine-scale species co-occurrence patterns in an old-growth temperate forest.Forest Ecology and Management,2009,257(10):2115- 2120.
[18] Stone L,Roberts A.The checkerboard score and species distributions.Oecologia,1990,85(1):74- 79.
[19] López R P,Valdivia S,Rivera M L,Rios R S.Co-occurrence patterns along a regional aridity gradient of the subtropical Andes do not support stress gradient hypotheses.PLoS One,2013,8(3):e58518.
[20] Gotelli N J,Ulrich W.The empirical bayes approach as a tool to identify non-random species associations.Oecologia,2010,162(2):463- 477.
[21] Veech J A,Miguel Araújo.The pairwise approach to analysing species co-occurrence.Journal of Biogeography,2014,41(6):1029-1035
[22] 陈蕾伊,沈海花,方精云.灌丛化草原:一种新的植被景观.自然杂志,2014,36(6):391- 396.
[23] Xu Z W,Wan S Q,Ren H Y,Han X G,Li M H,Cheng W X,Jiang Y.Effects of water and nitrogen addition on species turnover in temperate grasslands in northern China.PLoS One,2012,7(6):e39762.
[24] 辛小娟.氮、磷添加对亚高山草甸地上/地下生物量分配及植物功能群组成的影响[D].兰州:兰州大学,2011.
[25] 陈慧敏,石福习,杨桂生,张新厚,毛瑢.养分添加对三江平原沼泽化草甸植物群落组成和地上生物量的影响.生态学杂志,2016,35(6):1440- 1446.
[26] Blaser W J,Sitters J,Hart S P,Edwards P J,Olde Venterink H.Facilitative or competitive effects of woody plants on understorey vegetation depend on N-fixation,canopy shape and rainfall.Journal of Ecology,2013,101(6):1598- 1603.
[27] Dohn J,Dembélé F,Karembé M,Moustakas A,Amévor K A,Hanan N P.Tree effects on grass growth in savannas:competition,facilitation and the stress-gradient hypothesis.Journal of Ecology,2013,101(1):202- 209.
[28] Howard K S C,Eldridge D J,Soliveres S.Positive effects of shrubs on plant species diversity do not change along a gradient in grazing pressure in an arid shrubland.Basic and Applied Ecology,2012,13(2):159- 168.
[29] Jeltsch F,Milton S J,Dean W R J,Van Rooyen N,Moloney K A.Modelling the impact of small-scale heterogeneities on tree-grass coexistence in Semi-Arid Savannas.Journal of Ecology,1998,86(5):780- 793.
[30] 彭海英,李小雁,童绍玉.干旱半干旱区草原灌丛化研究进展.草业学报,2014,23(2):313- 322.
[31] 高玉葆,任安芝,王巍,王金龙.科尔沁沙地黄柳再生枝与现存枝形态和光合特征的比较.生态学报,2002,22(10):1758- 1764.
[32] Zhang G F,Zhao W Z.Species-specific traits determine shrub-annual interactions during a growing season.Journal of Arid Land,2015,7(3):403- 413.
[33] Sfenthourakis S,Tzanatos E,Giokas S.Species co-occurrence:the case of congeneric species and a causal approach to patterns of species association.Global Ecology and Biogeography,2006,15(1):39- 49.
[34] Arita H T.Species co-occurrence analysis:pairwise versus matrix-level approaches.Global Ecology and Biogeography,2016,25(11):1397- 1400.
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