荒漠绿洲过渡带柽柳种群结构与空间格局动态
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摘要
在塔克拉玛干沙漠北缘荒漠绿洲过渡带设置1 hm2研究样地,应用相邻格子法进行每木调查,从径级结构、空间分布格局、生命表、存活曲线和时间序列等方面研究柽柳种群结构与空间格局动态及其形成机理。结果表明:荒漠绿洲过渡带柽柳群落结构简单、物种稀少,多为单优群落。柽柳种群径级与高度结构均呈纺锤型,属衰退型种群;存活数、期望寿命(ex)随径级增加而降低,存活曲线接近DeeveyⅡ型,小径级死亡率、消失率与高度级失稳率较高,未来小径级、大径级灌木数量分别随时间推移减少与显著增多,更新资源匮乏使种群走向衰败。柽柳种群空间格局总体上为聚集分布,幼苗、小径级灌木为聚集分布,中、大径级灌木则为随机分布;不同径级的空间格局随尺度变化明显不同,小径级与大径级灌木分别在<3 m、<13 m和17—36 m尺度内呈随机分布,其他尺度均呈聚集分布,而中等径级灌木在研究尺度内呈随机分布,这是其繁殖特性、种内竞争、生境异质性与生态适应的结果。小径级与中等径级灌木、小径级与大径级灌木空间关联性分别在≥9 m、7—47 m空间尺度上相互排斥,致使幼小径级生长发育受抑而影响种群稳定。种群多度分布与环境的多元统计分析表明土壤水分、氮素是限制柽柳种群生长、分布和造成种群更新困难、衰退的主要因子。
Tamarix ramosissima is perennial shrub with strong resistance to drought,salinity,and wind. As one of the ecological keystone species of the Tarim Desert natural ecosystem,it plays a dominant role in maintaining the stabilized structure and function of the desert ecosystem and protecting the ecological safety of the oasis. To illuminate the ecological characteristics of a T. ramosissima population and to better understand the effects of biological and environmental factors on its population distribution,we studied the population structure,life table,survivorship curve,quantitative dynamics,and spatial distribution patterns of a T. ramosissima population at the desert-oasis ecotone of the Tarim Basin,Xinjiang,China.Ensuring the wise use,conservation,and preservation of the ecological bushwood in the Tarim Basin was the goal of this study. The age structure,life table,survivorship curve,and time sequence prediction of the T. ramosissima population were determined to analyze its population structure and dynamics. A simple community structure and a sparse species composition were found in the T. ramosissima at the desert-oasis ecotone of Tarim Basin. The age structure of the T. ramosissima population had a spindle-type shape with a low percentage of younger individuals,indicating that the population is likely to decline continuously. The number of surviving individuals and life expectancy of the T. ramosissima populationmonotonically decreased with increasing size class. The survivorship curve generally matched with Deevey II type. Small size shrubs are always accompanied with higher mortality rates and hazard rates,while highness class structure indicated low shrubs had higher lost stable rate. Seedlings could not be regenerated in time. The number of large size shrubs increased significantly with community development,which accelerates the shrunk of population size. This species generally had a clumped distribution pattern. Nevertheless,this pattern gradually changed over time from a clumped to a random pattern as shrubs matured from young to middle-aged to old-growth shrubs. Spatial variations of different growth stages changed obviously with different scales. Small size shrubs and large size shrubs had random distributions at scales of 0—3 m,0—13m and 17—36 m,respectively. They showed aggregated distributions at other scales. Mid-sized shrubs had a random distribution at all scales,which was related to habitat heterogeneity,reproductive characteristics,intraspecific competition,and ecological adaptation. Results of the spatial association analysis showed that small size shrubs were significantly negatively associated( mutually exclusive) to mid-size shrubs at scales ≥ 9 m,to large size shrubs at scales of 7—47 m,which were not conducive to the growth and regeneration of small individuals,further to affect population stability.Correlation and stepwise regression analysis between environmental factors and abundance distribution showed that the soil water and nitrogen contents were the main factors that restricted the growth,distribution,and regeneration.
Tamarix ramosissima is perennial shrub with strong resistance to drought,salinity,and wind. As one of the ecological keystone species of the Tarim Desert natural ecosystem,it plays a dominant role in maintaining the stabilized structure and function of the desert ecosystem and protecting the ecological safety of the oasis. To illuminate the ecological characteristics of a T. ramosissima population and to better understand the effects of biological and environmental factors on its population distribution,we studied the population structure,life table,survivorship curve,quantitative dynamics,and spatial distribution patterns of a T. ramosissima population at the desert-oasis ecotone of the Tarim Basin,Xinjiang,China.Ensuring the wise use,conservation,and preservation of the ecological bushwood in the Tarim Basin was the goal of this study. The age structure,life table,survivorship curve,and time sequence prediction of the T. ramosissima population were determined to analyze its population structure and dynamics. A simple community structure and a sparse species composition were found in the T. ramosissima at the desert-oasis ecotone of Tarim Basin. The age structure of the T. ramosissima population had a spindle-type shape with a low percentage of younger individuals,indicating that the population is likely to decline continuously. The number of surviving individuals and life expectancy of the T. ramosissima populationmonotonically decreased with increasing size class. The survivorship curve generally matched with Deevey II type. Small size shrubs are always accompanied with higher mortality rates and hazard rates,while highness class structure indicated low shrubs had higher lost stable rate. Seedlings could not be regenerated in time. The number of large size shrubs increased significantly with community development,which accelerates the shrunk of population size. This species generally had a clumped distribution pattern. Nevertheless,this pattern gradually changed over time from a clumped to a random pattern as shrubs matured from young to middle-aged to old-growth shrubs. Spatial variations of different growth stages changed obviously with different scales. Small size shrubs and large size shrubs had random distributions at scales of 0—3 m,0—13m and 17—36 m,respectively. They showed aggregated distributions at other scales. Mid-sized shrubs had a random distribution at all scales,which was related to habitat heterogeneity,reproductive characteristics,intraspecific competition,and ecological adaptation. Results of the spatial association analysis showed that small size shrubs were significantly negatively associated( mutually exclusive) to mid-size shrubs at scales ≥ 9 m,to large size shrubs at scales of 7—47 m,which were not conducive to the growth and regeneration of small individuals,further to affect population stability.Correlation and stepwise regression analysis between environmental factors and abundance distribution showed that the soil water and nitrogen contents were the main factors that restricted the growth,distribution,and regeneration.
引文
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[14]肖宜安,何平,李晓红,邓洪平.濒危植物长柄双花木自然种群数量动态.植物生态学报,2004,28(2):252-257.
[15]周洪华,陈亚宁,李卫红.塔里木河下游绿洲-荒漠过渡带植物多样性特征及优势种群分布格局.中国沙漠,2009,29(4):688-696.
[16]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.
[17]于军,王海珍,陈加利,韩路.塔里木河流域荒漠河岸林胡杨群落的空间格局研究.中国沙漠,2011,31(4):913-918.
[18]张春雨,赵秀海,王新怡,侯继华.长白山自然保护区红松阔叶林空间格局研究.北京林业大学学报,2006,28(增刊2):45-51.
[19] Rosner B. Fundamentals of Biostatistics. Sun S G,trans. 5th ed. Beijing:Science Press,2004:412-437.
[20]尹林克.柽柳属植物的生态适应性与引种.干旱区研究,2002,19(3):12-16.
[21]杨华,李艳丽,沈林,亢新刚.长白山云冷杉针阔混交林主要树种空间分布及其关联性.生态学报,2014,34(16):4698-4706.
[22] He F L,Legendre P,La Frankie J V. Distribution patterns of tree species in a Malaysian tropical rain forest. Journal of Vegetation Science,1997,8(1):105-114.
[23] Harms K E,Condit R,Hubbell S P,Foster R B. Habitat associations of trees and shrubs in a 50-ha neotropical forest plot. Journal of Ecology,2001,89(6):947-959.
[24] Lin Y C,Chang L W,Yang K C,Wang H H,Sun I F. Point patterns of tree distribution determined by habitat heterogeneity and dispersal limitation. Oecologia,2011,165(1):175-184.
[25] Condit R,Ashton P S,Baker P,Bunyavejchewin S,Gunatilleke S,Gunatilleke N,Hubbell S P,Foster R B,Itoh A,La Frankie J V,Lee H S,Losos E,Manokaran N,Sukumar R,Yamakura T. Spatial patterns in the distribution of tropical tree species. Science,2000,288(5470):1414-1418.
[26]赵峰侠,尹林克.荒漠内陆河岸胡杨和多枝柽柳幼苗种群空间分布格局及种间关联性.生态学杂志,2007,26(7):972-977.
[27]袁国富,张佩,薛沙沙,庄伟.沙丘多枝柽柳灌丛根层土壤含水量变化特征与根系水力提升证据.植物生态学报,2012,36(10):1033-1042.
[28] Wiegand T,Gunatilleke S,Gunatilleke N,Okuda T. Analyzing the spatial structure of a Sri Lankan tree species with multiple scales of clustering.Ecology,2007,88(12):3088-3102.
[29]刘普幸,张克新,霍华丽,潘竟虎.疏勒河中下游绿洲胡杨林土壤水盐的空间变化特征与成因.自然资源学报,2012,27(6):942-952.
[30] Glenn E,Tanner R,Mendez S,Kehret T,Moore D,Garcia J,Valdes C. Growth rates,salt tolerance and water use characteristics of native and invasive riparian plants from the delta of the Colorado River,Mexico. Journal of Arid Environment,1998,40(3):281-294.
[31]方敏彦,章明,丁品,张德罡.疏勒河中游高盐生境长穗柽柳群落格局研究.草业学报,2009,18(3):20-27.
[2]刘亚琦,刘加珍,陈永金,靖淑慧,冯若昂,毛甘霖.黄河三角洲潮间带柽柳灌丛的格局及结构动态研究.生态科学,2017,36(1):153-158.
[3]夏江宝,赵西梅,刘俊华,赵自国,刘庆,陈印平.黄河三角洲莱州湾湿地柽柳种群分布特征及其影响因素.生态学报,2016,36(15):4801-4808.
[4]郑姚闽,崔国发,雷霆,吴三雄,孙志成,袁海峰.甘肃敦煌西湖多枝柽柳群落特征和种群格局.北京林业大学学报,2010,32(4):34-44.
[5]王子丰,张淑荣,徐宗学,岳为峰,王国强,徐琛,赵捷.黑河中游山前荒漠和绿洲荒漠过渡带土壤植被空间分布特征研究.北京师范大学学报:自然科学版,2016,52(3):340-349.
[6]方欧娅,贾恒锋,邱红岩,任海保.青海省同德县乔木状甘蒙柽柳的年龄及其生长对环境的响应.植物生态学报,2017,41(7):738-748.
[7]刘华,董玲,李丕军,王文月,贾瑞琪.塔河流域不同生境条件下柽柳种群结构异质性特征.东北林业大学学报,2015,43(7):23-26,62-62.
[8]张佩,袁国富,庄伟,薛沙沙.黑河中游荒漠绿洲过渡带多枝柽柳对地下水位变化的生理生态响应与适应.生态学报,2011,31(22):6677-6687.
[9]徐高兴,王立,徐先英,金红喜,柴成武.民勤绿洲边缘地下水埋深对柽柳灌丛生长及物种多样性的影响.草原与草坪,2017,37(2):49-56.
[10]刘进辉,王雪芹,马洋.沙漠绿洲过渡带柽柳灌丛沙堆-丘间地系统土壤养分空间异质性.生态学报,2016,36(4):979-990.
[11]杨尧军,袁宏波,刘淑娟,张锦春,唐进年,丁峰.库姆塔格沙漠沙生柽柳种群与环境因子分析.干旱区研究,2013,30(5):827-831.
[12]程伟,吴宁,罗鹏.岷江上游林线附近岷江冷杉种群的生存分析.植物生态学报,2005,29(3):349-353.
[13]陈晓德,李旭光,王金锡.绵阳官司河流域长江防护林的群落高度级结构分析.植物生态学报,1997,21(4):376-385.
[14]肖宜安,何平,李晓红,邓洪平.濒危植物长柄双花木自然种群数量动态.植物生态学报,2004,28(2):252-257.
[15]周洪华,陈亚宁,李卫红.塔里木河下游绿洲-荒漠过渡带植物多样性特征及优势种群分布格局.中国沙漠,2009,29(4):688-696.
[16]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.
[17]于军,王海珍,陈加利,韩路.塔里木河流域荒漠河岸林胡杨群落的空间格局研究.中国沙漠,2011,31(4):913-918.
[18]张春雨,赵秀海,王新怡,侯继华.长白山自然保护区红松阔叶林空间格局研究.北京林业大学学报,2006,28(增刊2):45-51.
[19] Rosner B. Fundamentals of Biostatistics. Sun S G,trans. 5th ed. Beijing:Science Press,2004:412-437.
[20]尹林克.柽柳属植物的生态适应性与引种.干旱区研究,2002,19(3):12-16.
[21]杨华,李艳丽,沈林,亢新刚.长白山云冷杉针阔混交林主要树种空间分布及其关联性.生态学报,2014,34(16):4698-4706.
[22] He F L,Legendre P,La Frankie J V. Distribution patterns of tree species in a Malaysian tropical rain forest. Journal of Vegetation Science,1997,8(1):105-114.
[23] Harms K E,Condit R,Hubbell S P,Foster R B. Habitat associations of trees and shrubs in a 50-ha neotropical forest plot. Journal of Ecology,2001,89(6):947-959.
[24] Lin Y C,Chang L W,Yang K C,Wang H H,Sun I F. Point patterns of tree distribution determined by habitat heterogeneity and dispersal limitation. Oecologia,2011,165(1):175-184.
[25] Condit R,Ashton P S,Baker P,Bunyavejchewin S,Gunatilleke S,Gunatilleke N,Hubbell S P,Foster R B,Itoh A,La Frankie J V,Lee H S,Losos E,Manokaran N,Sukumar R,Yamakura T. Spatial patterns in the distribution of tropical tree species. Science,2000,288(5470):1414-1418.
[26]赵峰侠,尹林克.荒漠内陆河岸胡杨和多枝柽柳幼苗种群空间分布格局及种间关联性.生态学杂志,2007,26(7):972-977.
[27]袁国富,张佩,薛沙沙,庄伟.沙丘多枝柽柳灌丛根层土壤含水量变化特征与根系水力提升证据.植物生态学报,2012,36(10):1033-1042.
[28] Wiegand T,Gunatilleke S,Gunatilleke N,Okuda T. Analyzing the spatial structure of a Sri Lankan tree species with multiple scales of clustering.Ecology,2007,88(12):3088-3102.
[29]刘普幸,张克新,霍华丽,潘竟虎.疏勒河中下游绿洲胡杨林土壤水盐的空间变化特征与成因.自然资源学报,2012,27(6):942-952.
[30] Glenn E,Tanner R,Mendez S,Kehret T,Moore D,Garcia J,Valdes C. Growth rates,salt tolerance and water use characteristics of native and invasive riparian plants from the delta of the Colorado River,Mexico. Journal of Arid Environment,1998,40(3):281-294.
[31]方敏彦,章明,丁品,张德罡.疏勒河中游高盐生境长穗柽柳群落格局研究.草业学报,2009,18(3):20-27.