濒危植物树枫杜鹃(Rhododendron changii)种群结构及伴生群落特征
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摘要
分析了濒危植物树枫杜鹃种群结构及伴生群落物种组成、乔灌层物种多样性、垂直成层及个体径级结构等特征,并通过冗余分析(RDA)、典范对应分析(CCA)方法检验了树枫杜鹃种群及群落乔灌层物种组成与环境因子间的关系。结果表明:(1)树枫杜鹃种群径级结构呈单峰型,更新不连续,缺乏幼苗,幼株时期为又一死亡高峰期,种群呈衰退趋势;(2) 4个伴生群落中共有木本植物56种,隶属于23科40属,优势科为壳斗科、蔷薇科、槭树科、杜鹃花科,林下多箭竹(Fargesia spathacea);伴生群落的乔灌层多样性指数均较低,草本层不发达;群落1与群落2为高度1～1.5 m的灌丛,群落3为高度3～4 m的常绿阔叶-苔藓矮林,群落4为高度8～10 m的常绿落叶阔叶混交林;树枫杜鹃均位于群落下层,相比于乔木林群落,其在灌丛群落中生长较占优势;群落中耐阴种的小径级个体比例大,均呈增长趋势,林下竞争将持续增强;(3)前向选择及蒙特卡罗(Monte Carlo)检验表明,坡位对乔木层物种组成影响极显著,郁闭度与坡向对灌木层物种组成影响极显著(P<0.01);而枯落物厚度与空气相对湿度是影响树枫杜鹃幼苗的主要环境因子,林冠郁闭度、坡向则是影响幼株与成株的主要环境因子;同时,幼株与成株也分别受到林下物种密度与林下竹类覆盖度的影响(P<0.01)。因此,疏伐上层林木,人为干预林下幼树及箭竹的生长,并清理地表枯落物,对缓解树枫杜鹃濒危的现状及维持其种群的发展具有重要意义。
Rhododendron changii is a threatened plant species. Here,we investigated its population structure,species composition of companion communities,vertical stratification and plant diameter class,and species diversity in tree and shrub layers. Furthermore,the relationships between R. changii population and species composition of tree and shrub layers in community and environmental factors were examined using redundancy analysis( RDA) and canonical correspondence analysis( CCA). The results showed that:( 1) The diameter class pattern of R. changii was unimodal,indicating that the population renewability was limited in a discrete way and lack of seedlings. In addition,the sapling period was another peak of death,implying the population was continuing to decline.( 2) There were four companion communities around the population,containing a total of 56 woody species belonging to 23 families and 40 genera. The dominant families were Fagaceae,Rosaceae,Aceraceae,and Ericaceae,with a large number of Fargesia spathacea in the understory. The diversity index of tree and shrub layers was relatively low with an underdeveloped herbaceous layer. The companion communities 1 and 2 belonged to shrubs with an average height of 1-1.5 m,the community 3 was an evergreen broadleaf-mossy dwarf forest with an average height of 3-4 m,and the community 4 was an evergreen deciduous mixed forest with an average height of 8-10 m. R. changii grew in the lower layer in all companion communities,while dominated in shrub communities compared to arbor one. There was a large proportion of small-diametric individuals of shade-tolerant species in the community,indicating increasing competition in the understory.( 3) The results of the forward selection and Monte Carlo analysis showed that slope position exerted a significant impact on species composition of tree layer,while the canopy closeness and slope exposure controlled that of the shrub layer( P < 0. 01). Litter thickness and air humidity were two primary environmental factors affecting R. changii seedlings,while canopy coverage and slope exposure influenced the saplings and adults of R. changii.Besides,saplings and adult plants were also affected by the species density and bamboo coverage( P<0.01). In conclusion,it is important to decrease canopy cover,interfere with the growth of the other saplings and F. spathacea in forests and clean up litter layer,to relieve the endangered situation and maintain population development of R. changii.
Rhododendron changii is a threatened plant species. Here,we investigated its population structure,species composition of companion communities,vertical stratification and plant diameter class,and species diversity in tree and shrub layers. Furthermore,the relationships between R. changii population and species composition of tree and shrub layers in community and environmental factors were examined using redundancy analysis( RDA) and canonical correspondence analysis( CCA). The results showed that:( 1) The diameter class pattern of R. changii was unimodal,indicating that the population renewability was limited in a discrete way and lack of seedlings. In addition,the sapling period was another peak of death,implying the population was continuing to decline.( 2) There were four companion communities around the population,containing a total of 56 woody species belonging to 23 families and 40 genera. The dominant families were Fagaceae,Rosaceae,Aceraceae,and Ericaceae,with a large number of Fargesia spathacea in the understory. The diversity index of tree and shrub layers was relatively low with an underdeveloped herbaceous layer. The companion communities 1 and 2 belonged to shrubs with an average height of 1-1.5 m,the community 3 was an evergreen broadleaf-mossy dwarf forest with an average height of 3-4 m,and the community 4 was an evergreen deciduous mixed forest with an average height of 8-10 m. R. changii grew in the lower layer in all companion communities,while dominated in shrub communities compared to arbor one. There was a large proportion of small-diametric individuals of shade-tolerant species in the community,indicating increasing competition in the understory.( 3) The results of the forward selection and Monte Carlo analysis showed that slope position exerted a significant impact on species composition of tree layer,while the canopy closeness and slope exposure controlled that of the shrub layer( P < 0. 01). Litter thickness and air humidity were two primary environmental factors affecting R. changii seedlings,while canopy coverage and slope exposure influenced the saplings and adults of R. changii.Besides,saplings and adult plants were also affected by the species density and bamboo coverage( P<0.01). In conclusion,it is important to decrease canopy cover,interfere with the growth of the other saplings and F. spathacea in forests and clean up litter layer,to relieve the endangered situation and maintain population development of R. changii.
引文
鲍士旦.2000.土壤农化分析(第三版).北京:中国农业出版社.
柴胜丰,庄雪影,韦霄,等.2013.光照强度对濒危植物毛瓣金花茶光合生理特性的影响.西北植物学报,33(3):547-554.
达良俊,杨永川,宋永昌.2004.浙江天童国家森林公园常绿阔叶林主要组成种的种群结构及更新类型.植物生态学报,28(3):376-384.
符婵娟,刘艳红,赵本元.2009.神农架巴山冷杉群落更新特点及影响因素.生态学报,29(8):4179-4186.
耿玉英.2014.中国杜鹃花属植物.上海:上海科学技术出版社.
管康林,吴家森,范义荣,等.2001.华顶山云锦杜鹃林衰退原因及对策.浙江农林大学学报,18(2):195-197.
黄永涛,姚兰,艾训儒,等.2015.鄂西南两个自然保护区亚热带常绿落叶阔叶混交林类型及其常绿和落叶物种组成结构分析.植物生态学报,39(10):990-1002.
金慧,赵莹,尹航,等.2017.长白山濒危植物牛皮杜鹃(Rhododendron chrysanthum)种群数量特征与动态分析.生态学杂志,36(11):3123-3130.
李典谟,徐汝梅,马祖飞.2005.物种濒危机制和保育原理.北京:科学出版社.
李家湘,张旭,谢宗强,等.2015.湖南大围山杜鹃灌丛的群落组成及结构特征.生物多样性,23(6):815-823.
李俊清.2006.森林生态学.北京:高等教育出版社.
李晓笑,王清春,崔国发,等.2011.濒危植物梵净山冷杉野生种群结构及动态特征.西北植物学报,31(7):1479-1486.
刘振华,姚娜,杨凯,等.2012.树枫杜鹃菌根真菌分离与鉴定.林业科学研究,25(6):795-797.
蔺菲,郝占庆,李步杭,等.2007.长白山暗针叶林苔藓植物群落特征与林木更新的关系.生态学报,27(4):1308-1314.
蔺菲,郝占庆,叶吉.2006.苔藓植物对植物天然更新的影响.生态学杂志,25(4):456-460.
唐敏.2004.稀有植物太行花的保护生物学研究(博士学位论文).北京:中国科学院植物研究所.
王世雄,王孝安,李国庆,等.2010.陕西子午岭植物群落演替过程中物种多样性变化与环境解释.生态学报,30(6):1638-1647.
王小龙.2017.牛皮杜鹃生态适应能力研究(博士学位论文).长春:吉林大学.
王鑫,张华雨,李宗峰,等.2016.濒危植物崖柏群落特征及种群更新研究.北京林业大学学报,38(10):28-37.
熊高明,谢宗强,熊小刚,等.2003.神农架南坡珍稀植物独花兰的物候、繁殖及分布的群落特征.生态学报,23(1):173-179.
谢宗强,陈伟烈,路鹏,等.1999.濒危植物银杉的种群统计与年龄结构.生态学报,19(4):523-528.
颜超,王中生,安树青,等.2008.濒危植物银缕梅(Parrotia subaequalis)不同径级个体的光合能力差异与更新限制.生态学报,28(9):4153-4161.
晏海.2014.城市公园绿地小气候环境效应及其影响因子研究(博士学位论文).北京:北京林业大学.
杨旭,杨志玲,雷虓,等.2013.濒危植物凹叶厚朴幼苗更新及环境解释.林业科学,49(12):36-42.
张金屯.2011.数量生态学.北京:科学出版社.
张敬丽,吴雅文,吴红芝,等.2012.云南4种常绿杜鹃亚属植物种子萌发特性研究初探.云南农业大学学报:自然科学版,27(6):875-881.
张腾达.2014.金佛山林下小径竹类对乔木幼苗更新的影响(硕士学位论文).重庆:西南大学.
张文辉,祖元刚,刘国彬.2002.十种濒危植物的种群生态学特征及致危因素分析.生态学报,22(9):1512-1520.
张宇阳,马文宝,于涛,等.2018.梓叶槭的种群结构和群落特征.应用与环境生物学报,24(4):697-703.
周艳,陈训,韦小丽,等.2015.凋落物对迷人杜鹃幼苗更新和种子萌发的影响.林业科学,51(3):65-74.
周国宁,卜昭晖,陈绍云.1992.白花杜鹃生长发育规律与环境因子关系的研究.园艺学报,19(4):363-366.
朱建华,侯振宏,张治军,等.2007.气候变化与森林生态系统:影响、脆弱性与适应性.林业科学,43(11):138-145.
臧润国,董鸣,李俊清,等.2016.典型极小种群野生植物保护与恢复技术研究.生态学报,36(22):7130-7135.
Fabbro T,Krner C.2004.Altitudinal differences in flower traits and reproductive allocation.Flora,199:70-81.
Kameyama Y,Nakagoshi N,Nehira K.1999.Safe site for seedlings of Rhododendron metternichii var.hondoense.Plant Species Biology,14:237-242.
Mckinney ML.1997.Extinction vulnerability and selectivity:Combining ecological and paleontological views.Annual Review of Ecology&Systematics,28:495-516.
Médail F,Verlaque R.1997.Ecological characteristics and rarity of endemic plants from southeast France and Corsica:Implications for biodiversity conservation.Biological Conservation,80:269-281.
Root TL,Price JT,Hall KR,et al.2003.Fingerprints of global warming on wild animals and plants.Nature,421:57-60.
Tewksbury JJ,Huey RB,Deutsch CA.2008.Putting the Heat on Tropical Animals.Science,320:1296-1297.
柴胜丰,庄雪影,韦霄,等.2013.光照强度对濒危植物毛瓣金花茶光合生理特性的影响.西北植物学报,33(3):547-554.
达良俊,杨永川,宋永昌.2004.浙江天童国家森林公园常绿阔叶林主要组成种的种群结构及更新类型.植物生态学报,28(3):376-384.
符婵娟,刘艳红,赵本元.2009.神农架巴山冷杉群落更新特点及影响因素.生态学报,29(8):4179-4186.
耿玉英.2014.中国杜鹃花属植物.上海:上海科学技术出版社.
管康林,吴家森,范义荣,等.2001.华顶山云锦杜鹃林衰退原因及对策.浙江农林大学学报,18(2):195-197.
黄永涛,姚兰,艾训儒,等.2015.鄂西南两个自然保护区亚热带常绿落叶阔叶混交林类型及其常绿和落叶物种组成结构分析.植物生态学报,39(10):990-1002.
金慧,赵莹,尹航,等.2017.长白山濒危植物牛皮杜鹃(Rhododendron chrysanthum)种群数量特征与动态分析.生态学杂志,36(11):3123-3130.
李典谟,徐汝梅,马祖飞.2005.物种濒危机制和保育原理.北京:科学出版社.
李家湘,张旭,谢宗强,等.2015.湖南大围山杜鹃灌丛的群落组成及结构特征.生物多样性,23(6):815-823.
李俊清.2006.森林生态学.北京:高等教育出版社.
李晓笑,王清春,崔国发,等.2011.濒危植物梵净山冷杉野生种群结构及动态特征.西北植物学报,31(7):1479-1486.
刘振华,姚娜,杨凯,等.2012.树枫杜鹃菌根真菌分离与鉴定.林业科学研究,25(6):795-797.
蔺菲,郝占庆,李步杭,等.2007.长白山暗针叶林苔藓植物群落特征与林木更新的关系.生态学报,27(4):1308-1314.
蔺菲,郝占庆,叶吉.2006.苔藓植物对植物天然更新的影响.生态学杂志,25(4):456-460.
唐敏.2004.稀有植物太行花的保护生物学研究(博士学位论文).北京:中国科学院植物研究所.
王世雄,王孝安,李国庆,等.2010.陕西子午岭植物群落演替过程中物种多样性变化与环境解释.生态学报,30(6):1638-1647.
王小龙.2017.牛皮杜鹃生态适应能力研究(博士学位论文).长春:吉林大学.
王鑫,张华雨,李宗峰,等.2016.濒危植物崖柏群落特征及种群更新研究.北京林业大学学报,38(10):28-37.
熊高明,谢宗强,熊小刚,等.2003.神农架南坡珍稀植物独花兰的物候、繁殖及分布的群落特征.生态学报,23(1):173-179.
谢宗强,陈伟烈,路鹏,等.1999.濒危植物银杉的种群统计与年龄结构.生态学报,19(4):523-528.
颜超,王中生,安树青,等.2008.濒危植物银缕梅(Parrotia subaequalis)不同径级个体的光合能力差异与更新限制.生态学报,28(9):4153-4161.
晏海.2014.城市公园绿地小气候环境效应及其影响因子研究(博士学位论文).北京:北京林业大学.
杨旭,杨志玲,雷虓,等.2013.濒危植物凹叶厚朴幼苗更新及环境解释.林业科学,49(12):36-42.
张金屯.2011.数量生态学.北京:科学出版社.
张敬丽,吴雅文,吴红芝,等.2012.云南4种常绿杜鹃亚属植物种子萌发特性研究初探.云南农业大学学报:自然科学版,27(6):875-881.
张腾达.2014.金佛山林下小径竹类对乔木幼苗更新的影响(硕士学位论文).重庆:西南大学.
张文辉,祖元刚,刘国彬.2002.十种濒危植物的种群生态学特征及致危因素分析.生态学报,22(9):1512-1520.
张宇阳,马文宝,于涛,等.2018.梓叶槭的种群结构和群落特征.应用与环境生物学报,24(4):697-703.
周艳,陈训,韦小丽,等.2015.凋落物对迷人杜鹃幼苗更新和种子萌发的影响.林业科学,51(3):65-74.
周国宁,卜昭晖,陈绍云.1992.白花杜鹃生长发育规律与环境因子关系的研究.园艺学报,19(4):363-366.
朱建华,侯振宏,张治军,等.2007.气候变化与森林生态系统:影响、脆弱性与适应性.林业科学,43(11):138-145.
臧润国,董鸣,李俊清,等.2016.典型极小种群野生植物保护与恢复技术研究.生态学报,36(22):7130-7135.
Fabbro T,Krner C.2004.Altitudinal differences in flower traits and reproductive allocation.Flora,199:70-81.
Kameyama Y,Nakagoshi N,Nehira K.1999.Safe site for seedlings of Rhododendron metternichii var.hondoense.Plant Species Biology,14:237-242.
Mckinney ML.1997.Extinction vulnerability and selectivity:Combining ecological and paleontological views.Annual Review of Ecology&Systematics,28:495-516.
Médail F,Verlaque R.1997.Ecological characteristics and rarity of endemic plants from southeast France and Corsica:Implications for biodiversity conservation.Biological Conservation,80:269-281.
Root TL,Price JT,Hall KR,et al.2003.Fingerprints of global warming on wild animals and plants.Nature,421:57-60.
Tewksbury JJ,Huey RB,Deutsch CA.2008.Putting the Heat on Tropical Animals.Science,320:1296-1297.