热带海岸香蒲桃天然次生林群落优势种群种间联结性
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摘要
【目的】研究热带海岸香蒲桃天然次生林群落乔木层、灌木层、草本层各优势种群的种间关联性,以期阐明群落演替与种对间关联性的内在规律,为海岸香蒲桃天然次生林的保护、恢复与重建提供理论依据。【方法】选取海南文昌热带海岸典型香蒲桃天然次生林群落,采用方差比率法测定乔木层、灌木层、草本层优势种群的总体关联性,使用χ~2检验2个种群关联与否,进一步用Ochiai指数来表征关联程度。【结果】乔木层的总体关联性表现出不显著正关联,正关联种对数为54对,负关联种对数为49对,无关联种对数为17对,分别占总对数的45.0%,40.8%和14.2%;香蒲桃与群落中其他乔木树种无关联,具有较强的独立性;灌木层负关联种对数有33对,占总对数的43.6%,这些负关联的树种多以阳性植物组成为主;竹叶木姜子和九节易与其他物种形成正关联种对,亦是灌木层互利共生的主要树种;草本层的总体关联性为显著负关联(P<0.05),正关联种对数较少(12%),负关联种对数比例较高(78%),群落处于不稳定状态,容易受外界干扰而演替;海金沙和铺地黍这2物种在草本层中互利共生。【结论】热带海岸香蒲桃天然次生林群落优势种群种对间种间关联明显,物种之间存在一定的独立性,整体趋向稳定发展。实践中为加快热带海岸次生林演替进程,应以正联结性较强的树种对(山楝-黄槿、竹叶木姜子-九节、东风桔-露兜树)作为海防林的造林树种。
【Objective】Interspecific associations of dominant plant populations respectively in the layers of tree,shrub and herb in communities of secondary natural forest of Syzygium odoratum in tropical coast were studied to reveal the inherent linkages of interactions among different populations during the process of community development,and to clarify the natural rules of community development in relation to the succession and interspecific association,as well as to provide a theoretical basis for the recovery of coastal forests in China.【Method】Typical communities of secondary natural forest of S.odoratum in Hainan Island were selected for the study.2 × 2 contingency tables of species presence/absence data were used as the basis for the analysis of overall interspecific associations among dominant plant populations in different layers of tree,shrub and herb by using the variance ratio method.χ2 test and Ochiai index were used to characterize interspecific associations.【Results】Interspecific association in the tree layer was not significantly positive.For the 171 tree species,although there were collocation at some degrees,but a stable and harmonious collocation was not found.S.odoratum was not associated with other tree species.It was strongly independent.The inter-specific association of Casuarina equisetifolia and Pittosporum tobira were in significant positive correlation.The shrub layer displayed significant positive correlation in the inter-specific association (P < 0.05).Compared with the tree layer,inter-specific association in the herb layer was strongly negative.【Conclusion 】In the tropical coastal forest community,dominant species of displayed significant interspecific association and there existed a certain degree of independence between species and overall trend of stable development.In order to speed up the process of succession of the tropical coast secondary forest,the species withstrong coupling should be protected in practice.
【Objective】Interspecific associations of dominant plant populations respectively in the layers of tree,shrub and herb in communities of secondary natural forest of Syzygium odoratum in tropical coast were studied to reveal the inherent linkages of interactions among different populations during the process of community development,and to clarify the natural rules of community development in relation to the succession and interspecific association,as well as to provide a theoretical basis for the recovery of coastal forests in China.【Method】Typical communities of secondary natural forest of S.odoratum in Hainan Island were selected for the study.2 × 2 contingency tables of species presence/absence data were used as the basis for the analysis of overall interspecific associations among dominant plant populations in different layers of tree,shrub and herb by using the variance ratio method.χ2 test and Ochiai index were used to characterize interspecific associations.【Results】Interspecific association in the tree layer was not significantly positive.For the 171 tree species,although there were collocation at some degrees,but a stable and harmonious collocation was not found.S.odoratum was not associated with other tree species.It was strongly independent.The inter-specific association of Casuarina equisetifolia and Pittosporum tobira were in significant positive correlation.The shrub layer displayed significant positive correlation in the inter-specific association (P < 0.05).Compared with the tree layer,inter-specific association in the herb layer was strongly negative.【Conclusion 】In the tropical coastal forest community,dominant species of displayed significant interspecific association and there existed a certain degree of independence between species and overall trend of stable development.In order to speed up the process of succession of the tropical coast secondary forest,the species withstrong coupling should be protected in practice.
引文
陈玉凯,杨小波,李东海,等.2011.海南霸王岭海南油杉群落优势种群的种间联结性研究.植物科学学报,29(3):278-287.(Chen Y K,Yang X B,Li D H,et al.2011.The population of Hainan Bawangling advantage Hainan Keteleeria community study on the interspecific association.Plant Science Journal,29(3):278-287.[in Chinese])
程瑞梅,王瑞丽,刘泽彬,等.2013.三峡库区栲属群落主要乔木种群的种间联结性.林业科学,49(5):36-42.(Cheng R M,Wang R L,Liu Z B,et al.2013.The interspecific association of the main tree populations of Castanopsis community in Three Gorges Reservoir area.Scientia Silvae Sinicae,49(5):36-42.[in Chinese])
戈峰.2008.现代生态学.北京:科学出版社.(Ge F.2008.Modern ecology.Beijing:Science Press.[in Chinese])
郝建华,刘倩倩,强胜.2009.菊科入侵植物三叶鬼针草的繁殖特征及其与入侵性的关系.植物学报,44(6):656-665.(Hao J H,Liu Q Q,Qiang S.2009.Reproductive characteristics of Bidens pilosa and invasive.Journal of Plant Science,44(6):656-665.[in Chinese])
胡文强,黄世能,李家湘,等.2013.南岭石坑崆山顶矮林乔木优势种群的种间关联性.生态学杂志,32(10):2665-2671.(Hu W Q,Huang S N,Li J X,et al.2013.Population of Nanling Mountain Shikengkong elfin forest dominant tree species association.Chinese Journal of Ecology,32(10):2665-2671.[in Chinese])
黄彩霞,邓贤兰,汪维,等.2007.井冈山自然保护区南方红豆杉群落优势种种间联结性研究.江西林业科技,3:1-3+10.(Huang C X,Deng X L,Wang W,et al.2007.Study on the diversity of the community characteristics of Taxus mairei community in Jinggangshan nature reserve.Jiangxi Forestry Science and Technology,3:1-3+10.[in Chinese])
李佳灵,林育成,王旭,等.2013.海南吊罗山不同海拔高度热带雨林种间联结性对比研究.热带作物学报,34(3):584-590.(Li J L,Lin Y C,Wang X,et al.2013.Comparative study on interspecific association of tropical rain forests in Hainan,Diaoluoshan.Journal of Tropical Crops,34(3):584-590.[in Chinese])
刘珏宏,高慧,张丽红,等.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(9):1016-1024.(Liu J H,Gao H,Zhang L H,et al.2010.Interspecific association characteristics in Inner Mongolia Xilinguole steppe of Stipa grandisStipa krylovii community analysis.Chinese Journal of Plant Ecology,34(9):1016-1024.[in Chinese])
覃林.2009.生态统计学.北京:中国林业出版社:91-93.(Qin L.2009.Ecological statistics.Beijing:Chinese Forestry Press:91-93.[in Chinese])
奇凯,张春雨,侯继华,等.2010.赤峰市沙地油松林草本植物多样性及种间关联动态.生态学报,30(18):5106-5112.(Qi K,Zhang C Y,Hou J H,et al.2010.Studies on the diversity and interspecific association of herbaceous plant species of Pinus tabulaeformis forest in sandy land of Chifeng.Journal of Ecology,30(18):5106-5112.[in Chinese])
王伟,崔红,陈亮,等.2000.盐胁迫对不同生境铺地黍叶片蛋白质合成的影响.厦门大学学报:自然科学版,39(3):417-420.(Wang W,Cui H,Chen L,et al.2000.Effects of salt stress on leaf protein in different habitats Torpedograss synthesis.Journal of Xiamen University:Natural Science Edition,39(3):417-420.[in Chinese])
王文进,张明,刘福德,等.2007.海南岛吊罗山热带山地雨林两个演替阶段的种间联结性.生物多样性,15(3):257-263.(Wang W J,Zhang M,Liu F D,et al.2007.Interspecific association of two successional stages of tropical montane rain forest in Luoshan,Hainan Island,China.Bio-Diversity,15(3):257-263.[in Chinese])
杨青青,杨众养,薛杨,等.2015.海南文昌滨海台地不同森林林下灌草多样性及其与土壤因子关联分析.热带作物学报,36(12):2238-2244.(Yang Q Q,Yang Z Y,Xue Y,et al.2015.Diversity of shrub grass and its correlation with soil factors in Wenchang coastal platform of Hainan.Journal of Tropical Crops,36(12):2238-2244.[in Chinese])
杨琦,周婧,陶楚,等.2014.海南岛铜鼓岭热带常绿季雨矮林2个演替阶段的种间联结性对比研究.中国农学通报,30(22):8-15.(Yang Q,Zhou J,Tao C,et al.2014.Hainan Island Tongguling tropical evergreen monsoon elfin forest 2 successional stages of the interspecific association of the comparative study.China Agricultural Science Bulletin,30(22):8-15.[in Chinese])
杨沅志.2011.深圳市大鹏半岛海堤生物安全防护现状调查与评价.林业调查规划,36(01):63-68.(Yang Y Z.2011.Investigation and evaluation of current status of biological safety of Shenzhen Dapeng Peninsula.Forestry Investigation and Planning,36(01):63-68.[in Chinese])
喻本德,叶有华,郭微,等.2014.生态保护红线分区建设模式研究---以广东大鹏半岛为例.生态环境学报,23(6):962-971.(Yu B D,Ye Y H,Guo W,et al.2014.Research on the partition construction mode for ecological essential line:a case study in Dapeng Peninsula,Guangdong Province.Ecology and Environmental Sciences,23(6):962-971.[in Chinese])
张明霞,王得祥,康冰,等.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.(Zhang M X,Wang D X,Kang B,et al.2015.The interspecific association of dominant species in Qinling Mountains Huashan pine natural forest.Scientia Silvae Sinicae,51(1):12-21.[in Chinese])
张永夏,陈红锋,秦新生,等.2007.深圳大鹏半岛“风水林”香蒲桃群落特征及物种多样性研究.广西植物,27(4):596-603.(Zhang Y X,Chen H F,Qin X S,et al.2007.Shenzhen Dapeng Peninsula Syzygium odoratum community characteristics and species diversity of Syzygium odoratum Community in Dapeng Pninsula,Shenzhen.Plants in Guangxi,27(4):596-603.[in Chinese])
张悦,郭利平,易雪梅,等.2015.长白山北坡3个森林群落主要树种种间联结性.生态学报,35(1):106-115.(Zhang Y,Guo L P,Yi X M,et al.2015.Analysis of interspecific associations among major tree species in three forest communities on the north slope of Changbai Mountain.Journal of Ecology,35(1):106-115.[in Chinese])
Fisher R A.1943.The relation between the number of species and the number of individuals in a random sample of an animal population.Journal of Animal Ecology,12(1):42-58.
Forbes S A.1907.On the local distribution of certain Illinois fishes:an essay in statistical ecology.Bill,Illinois State Laboratory Natural History,7(8):237-303.
Gidudu B,Copeland R S,Wanda F,et al.2011.Distribution,interspecific associations and abundance of aquatic plants in Lake Bisina,Uganda.Journal of Aquatic Plant Management,49(1):19-27.
Green A J,Mustapha E H.2006.Interspecific associations in habitat use between marbled teal and other waterbirds wintering at Sidi Bou Ghaba,Morocco.Ardeola,53(1):99-106.
Lan G Y,Getzin S,Wiegand T,et al.2012.Spatial distribution and interspecific associations of tree species in a tropical seasonal rain forest of China.Plos One,7(9):1-9.
Li Y D,Xu H,Chen D X,et al.2008.Division of ecological species groups and functional groups based on interspecific association-a case study of the tree layer in the tropical lowland rainforest of Jianfenling in Hainan Island,China.Frontiers of Forestry in China,3(4):407-415.
Schluter D.1984.A variance test for detecting species associations,with some example applications.Ecology,65(3):998-1005.
Wang X G,Wiegand T,Hao Z,et al.2010.Species associations in an old-growth temperate forest in north-eastern China.Journal of Ecology,98:674-686.
Wiegand T,Gunatilleke S,Gunatilleke N.2007.Species associations in a heterogeneous Sri Lankan dipterocarp forest.American Naturalist,170:77-95.
Zhang Z H,Hu G,Zhu J D,et al.2010.Spatial patterns and interspecific associations of dominant tree species in two old growth karst forests,SW China.Ecological Research,25:1151-1160.
程瑞梅,王瑞丽,刘泽彬,等.2013.三峡库区栲属群落主要乔木种群的种间联结性.林业科学,49(5):36-42.(Cheng R M,Wang R L,Liu Z B,et al.2013.The interspecific association of the main tree populations of Castanopsis community in Three Gorges Reservoir area.Scientia Silvae Sinicae,49(5):36-42.[in Chinese])
戈峰.2008.现代生态学.北京:科学出版社.(Ge F.2008.Modern ecology.Beijing:Science Press.[in Chinese])
郝建华,刘倩倩,强胜.2009.菊科入侵植物三叶鬼针草的繁殖特征及其与入侵性的关系.植物学报,44(6):656-665.(Hao J H,Liu Q Q,Qiang S.2009.Reproductive characteristics of Bidens pilosa and invasive.Journal of Plant Science,44(6):656-665.[in Chinese])
胡文强,黄世能,李家湘,等.2013.南岭石坑崆山顶矮林乔木优势种群的种间关联性.生态学杂志,32(10):2665-2671.(Hu W Q,Huang S N,Li J X,et al.2013.Population of Nanling Mountain Shikengkong elfin forest dominant tree species association.Chinese Journal of Ecology,32(10):2665-2671.[in Chinese])
黄彩霞,邓贤兰,汪维,等.2007.井冈山自然保护区南方红豆杉群落优势种种间联结性研究.江西林业科技,3:1-3+10.(Huang C X,Deng X L,Wang W,et al.2007.Study on the diversity of the community characteristics of Taxus mairei community in Jinggangshan nature reserve.Jiangxi Forestry Science and Technology,3:1-3+10.[in Chinese])
李佳灵,林育成,王旭,等.2013.海南吊罗山不同海拔高度热带雨林种间联结性对比研究.热带作物学报,34(3):584-590.(Li J L,Lin Y C,Wang X,et al.2013.Comparative study on interspecific association of tropical rain forests in Hainan,Diaoluoshan.Journal of Tropical Crops,34(3):584-590.[in Chinese])
刘珏宏,高慧,张丽红,等.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(9):1016-1024.(Liu J H,Gao H,Zhang L H,et al.2010.Interspecific association characteristics in Inner Mongolia Xilinguole steppe of Stipa grandisStipa krylovii community analysis.Chinese Journal of Plant Ecology,34(9):1016-1024.[in Chinese])
覃林.2009.生态统计学.北京:中国林业出版社:91-93.(Qin L.2009.Ecological statistics.Beijing:Chinese Forestry Press:91-93.[in Chinese])
奇凯,张春雨,侯继华,等.2010.赤峰市沙地油松林草本植物多样性及种间关联动态.生态学报,30(18):5106-5112.(Qi K,Zhang C Y,Hou J H,et al.2010.Studies on the diversity and interspecific association of herbaceous plant species of Pinus tabulaeformis forest in sandy land of Chifeng.Journal of Ecology,30(18):5106-5112.[in Chinese])
王伟,崔红,陈亮,等.2000.盐胁迫对不同生境铺地黍叶片蛋白质合成的影响.厦门大学学报:自然科学版,39(3):417-420.(Wang W,Cui H,Chen L,et al.2000.Effects of salt stress on leaf protein in different habitats Torpedograss synthesis.Journal of Xiamen University:Natural Science Edition,39(3):417-420.[in Chinese])
王文进,张明,刘福德,等.2007.海南岛吊罗山热带山地雨林两个演替阶段的种间联结性.生物多样性,15(3):257-263.(Wang W J,Zhang M,Liu F D,et al.2007.Interspecific association of two successional stages of tropical montane rain forest in Luoshan,Hainan Island,China.Bio-Diversity,15(3):257-263.[in Chinese])
杨青青,杨众养,薛杨,等.2015.海南文昌滨海台地不同森林林下灌草多样性及其与土壤因子关联分析.热带作物学报,36(12):2238-2244.(Yang Q Q,Yang Z Y,Xue Y,et al.2015.Diversity of shrub grass and its correlation with soil factors in Wenchang coastal platform of Hainan.Journal of Tropical Crops,36(12):2238-2244.[in Chinese])
杨琦,周婧,陶楚,等.2014.海南岛铜鼓岭热带常绿季雨矮林2个演替阶段的种间联结性对比研究.中国农学通报,30(22):8-15.(Yang Q,Zhou J,Tao C,et al.2014.Hainan Island Tongguling tropical evergreen monsoon elfin forest 2 successional stages of the interspecific association of the comparative study.China Agricultural Science Bulletin,30(22):8-15.[in Chinese])
杨沅志.2011.深圳市大鹏半岛海堤生物安全防护现状调查与评价.林业调查规划,36(01):63-68.(Yang Y Z.2011.Investigation and evaluation of current status of biological safety of Shenzhen Dapeng Peninsula.Forestry Investigation and Planning,36(01):63-68.[in Chinese])
喻本德,叶有华,郭微,等.2014.生态保护红线分区建设模式研究---以广东大鹏半岛为例.生态环境学报,23(6):962-971.(Yu B D,Ye Y H,Guo W,et al.2014.Research on the partition construction mode for ecological essential line:a case study in Dapeng Peninsula,Guangdong Province.Ecology and Environmental Sciences,23(6):962-971.[in Chinese])
张明霞,王得祥,康冰,等.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.(Zhang M X,Wang D X,Kang B,et al.2015.The interspecific association of dominant species in Qinling Mountains Huashan pine natural forest.Scientia Silvae Sinicae,51(1):12-21.[in Chinese])
张永夏,陈红锋,秦新生,等.2007.深圳大鹏半岛“风水林”香蒲桃群落特征及物种多样性研究.广西植物,27(4):596-603.(Zhang Y X,Chen H F,Qin X S,et al.2007.Shenzhen Dapeng Peninsula Syzygium odoratum community characteristics and species diversity of Syzygium odoratum Community in Dapeng Pninsula,Shenzhen.Plants in Guangxi,27(4):596-603.[in Chinese])
张悦,郭利平,易雪梅,等.2015.长白山北坡3个森林群落主要树种种间联结性.生态学报,35(1):106-115.(Zhang Y,Guo L P,Yi X M,et al.2015.Analysis of interspecific associations among major tree species in three forest communities on the north slope of Changbai Mountain.Journal of Ecology,35(1):106-115.[in Chinese])
Fisher R A.1943.The relation between the number of species and the number of individuals in a random sample of an animal population.Journal of Animal Ecology,12(1):42-58.
Forbes S A.1907.On the local distribution of certain Illinois fishes:an essay in statistical ecology.Bill,Illinois State Laboratory Natural History,7(8):237-303.
Gidudu B,Copeland R S,Wanda F,et al.2011.Distribution,interspecific associations and abundance of aquatic plants in Lake Bisina,Uganda.Journal of Aquatic Plant Management,49(1):19-27.
Green A J,Mustapha E H.2006.Interspecific associations in habitat use between marbled teal and other waterbirds wintering at Sidi Bou Ghaba,Morocco.Ardeola,53(1):99-106.
Lan G Y,Getzin S,Wiegand T,et al.2012.Spatial distribution and interspecific associations of tree species in a tropical seasonal rain forest of China.Plos One,7(9):1-9.
Li Y D,Xu H,Chen D X,et al.2008.Division of ecological species groups and functional groups based on interspecific association-a case study of the tree layer in the tropical lowland rainforest of Jianfenling in Hainan Island,China.Frontiers of Forestry in China,3(4):407-415.
Schluter D.1984.A variance test for detecting species associations,with some example applications.Ecology,65(3):998-1005.
Wang X G,Wiegand T,Hao Z,et al.2010.Species associations in an old-growth temperate forest in north-eastern China.Journal of Ecology,98:674-686.
Wiegand T,Gunatilleke S,Gunatilleke N.2007.Species associations in a heterogeneous Sri Lankan dipterocarp forest.American Naturalist,170:77-95.
Zhang Z H,Hu G,Zhu J D,et al.2010.Spatial patterns and interspecific associations of dominant tree species in two old growth karst forests,SW China.Ecological Research,25:1151-1160.