东北阔叶红松林群落分类、排序及物种多样性比较
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摘要
植物群落是植物与环境相互作用的产物,探讨森林群落的分布格局、多样性变化规律及其与环境因子的关系,有助于该区森林群落稳定性的维持和生物多样性保护。在东北阔叶红松林区,沿纬度梯度在典型区域选择未受干扰的原始阔叶红松林进行群落调查,并运用多元回归树(MRT)、物种多样性指数比较和典范对应分析(CCA)方法,对森林群落进行分类、比较和排序。结果表明:东北阔叶红松林森林群落可分为4个类型,不同群落类型间物种组成及多样性差异显著。物种丰富度及多样性指数均为低纬度区的千金榆-枫桦-红松林显著高于高纬度地区的冷杉-红松林群落。CCA排序结果较好地反映了各群落类型的分布范围及其与环境因子的关系,其变化格局主要受温度和降水的影响,其次是土壤养分。该结果为气候变化下阔叶红松林的管理和保护提供了相应的理论依据。
The plant community is the product of the interaction between plants and the environment,and the interaction process is associated with changes in community structure and species diversity. Exploring the distribution of forest communities,species diversity patterns,and their relationship with environmental factors is important for protecting forest stability and biodiversity. Quantitative analysis is an important approach in research on community classification and sorting and can profoundly reveal the ecological relationship between plant communities and environmental factors. The study of species diversity patterns can reflect the ecological characteristics of plant communities and their adaptability to environment change. The Korean pine broadleaved forest(KBF) is the local climax vegetation and mainly distributed in northeast China.It has a complex forest structure and rich biodiversity and is very important in maintaining the regional biodiversity.Therefore,it is vital to conduct a case study regarding community classification,ordination,and species diversity patterns in broadleaved Korean pine forests. In recent years, many studies regarding KBF have been conducted from various perspectives,but forest community classification, sorting, and comparison research using quantitative ecology is still lacking. Along a latitudinal gradient,we investigated the forest community structure with 40 plots(20 m × 20 m). Then,forest community classification,ordination,and comparison of species diversity were conducted using the methods of MRT(multiple regression tree),CCA(correspondence analysis),and comparison of species diversity indexes. The results showed that the MRT classified the forest communities into four types by relating different aspects of species composition to environmental data:(I) Abies fabri-Pinus koraiensis Association,(II) Quercus mongolica-Acer mono-Pinus koraiensis Association,(III) Fraxinus mandshurica-Acer tegmentosum-Pinus koraiensis Association,(IV) Carpinus cordata-Betula costata-Pinus koraiensis Association. Species composition and diversity of forest communities differed greatly among the four types. The richness and species diversity index of Carpinus cordata-Betula costata-Pinus koraiensis Association were significantly higher than that of the Abies fabri Craib-Pinus koraiensis Association. The results of CCA were consistent with the forest community classification from MRT. It clearly reflected the distribution range of various community types and the relationships between community distribution and environmental factors. The Abies fabri-Pinus koraiensis Association corresponded to colder habitat,whereas the Carpinus cordata-Betula costata-Pinus koraiensis Association corresponded to warmer and wetter habitat. The Quercus mongolica-Acer mono-Pinus koraiensis Association tended to be in habitats with fertile soil. The distribution of the Fraxinus mandshurica-Acer tegmentosum-Pinus koraiensis Association had no special preference for environmental conditions. Temperature and precipitation were the main environmental factors affecting the distribution pattern of forest communities,whereas soil nutrients had smaller effects. These results help to more thoroughly understand the space distribution pattern and ecological characteristics of the forest community. They also provide the corresponding theoretical basis for the management and protection of broadleaved Korean pine forests in the future.
The plant community is the product of the interaction between plants and the environment,and the interaction process is associated with changes in community structure and species diversity. Exploring the distribution of forest communities,species diversity patterns,and their relationship with environmental factors is important for protecting forest stability and biodiversity. Quantitative analysis is an important approach in research on community classification and sorting and can profoundly reveal the ecological relationship between plant communities and environmental factors. The study of species diversity patterns can reflect the ecological characteristics of plant communities and their adaptability to environment change. The Korean pine broadleaved forest(KBF) is the local climax vegetation and mainly distributed in northeast China.It has a complex forest structure and rich biodiversity and is very important in maintaining the regional biodiversity.Therefore,it is vital to conduct a case study regarding community classification,ordination,and species diversity patterns in broadleaved Korean pine forests. In recent years, many studies regarding KBF have been conducted from various perspectives,but forest community classification, sorting, and comparison research using quantitative ecology is still lacking. Along a latitudinal gradient,we investigated the forest community structure with 40 plots(20 m × 20 m). Then,forest community classification,ordination,and comparison of species diversity were conducted using the methods of MRT(multiple regression tree),CCA(correspondence analysis),and comparison of species diversity indexes. The results showed that the MRT classified the forest communities into four types by relating different aspects of species composition to environmental data:(I) Abies fabri-Pinus koraiensis Association,(II) Quercus mongolica-Acer mono-Pinus koraiensis Association,(III) Fraxinus mandshurica-Acer tegmentosum-Pinus koraiensis Association,(IV) Carpinus cordata-Betula costata-Pinus koraiensis Association. Species composition and diversity of forest communities differed greatly among the four types. The richness and species diversity index of Carpinus cordata-Betula costata-Pinus koraiensis Association were significantly higher than that of the Abies fabri Craib-Pinus koraiensis Association. The results of CCA were consistent with the forest community classification from MRT. It clearly reflected the distribution range of various community types and the relationships between community distribution and environmental factors. The Abies fabri-Pinus koraiensis Association corresponded to colder habitat,whereas the Carpinus cordata-Betula costata-Pinus koraiensis Association corresponded to warmer and wetter habitat. The Quercus mongolica-Acer mono-Pinus koraiensis Association tended to be in habitats with fertile soil. The distribution of the Fraxinus mandshurica-Acer tegmentosum-Pinus koraiensis Association had no special preference for environmental conditions. Temperature and precipitation were the main environmental factors affecting the distribution pattern of forest communities,whereas soil nutrients had smaller effects. These results help to more thoroughly understand the space distribution pattern and ecological characteristics of the forest community. They also provide the corresponding theoretical basis for the management and protection of broadleaved Korean pine forests in the future.
引文
[1]王世雄,王孝安,李国庆,郭华,朱志红.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,2010,30(6):1638-1647.
[2]白晓航,张金屯,曹科,王云泉,Sadia S,曹格.河北小五台山国家级自然保护区森林群落与环境的关系.生态学报,2017,37(11):3683-3696.
[3]田永强,俞超超,王磊,黄邦钦.福建九龙江北溪浮游植物群落分布特征及其影响因子.应用生态学报,2012,23(9):2559-2565.
[4]武春华.黄土丘陵区典型植物群落特征及其与环境因子的关系[D].北京:中国科学院研究生院,2008.
[5]赵从举,康慕谊,雷加强.准噶尔盆地典型地段植物群落及其与环境因子的关系.生态学报,2011,31(10):2669-2677.
[6]袁铁象,张合平,欧芷阳,谭一波.地形对桂西南喀斯特山地森林地表植物多样性及分布格局的影响.应用生态学报,2014,25(10):2803-2810.
[7]任萍.黄土高原马栏林区植物群落物种多度研究[D].西安:陕西师范大学,2010.
[8]苏日古嘎,张金屯,张斌,程佳佳,田世广,张钦弟,刘素军.松山自然保护区森林群落的数量分类和排序.生态学报,2010,30(10):2621-2629.
[9]张文静,张钦弟,王晶,冯飞,毕润成.多元回归树与双向指示种分析在群落分类中的应用比较.植物生态学报,2015,39(6):586-592.
[10]张金屯.数量生态学.北京:科学出版社,2011:123-127.
[11] Virtanen R,Luoto M,RmT,Mikkola K,Hjort J,Grytnes J A,Birks H J B. Recent vegetation changes at the high-latitude tree line ecotone are controlled by geomorphological disturbance,productivity and diversity. Global Ecology and Biogeography,2010,19(6):810-821.
[12]李博.生态学.北京:高等教育出版社,2000:168-172.
[13]代力民,谷会岩,邵国凡,王庆礼.中国长白山阔叶红松林.沈阳:辽宁科学技术出版社,2004:5-11.
[14]吴正方,邓慧平.东北阔叶红松林全球气候变化响应研究.地理学报,1996,51(增刊):81-91.
[15]吴正方.东北阔叶红松林分布区生态气候适宜性及全球气候变化影响评价.应用生态学报,2003,14(5):771-775.
[16]贾翔,马芳芳,周旺明,周莉,于大炮,秦静,代力民.气候变化对阔叶红松林潜在地理分布区的影响.生态学报,2017,37(2):464-473.
[17]曹伟,李岩,王树良,于兴华,朱彩霞,刘童燕.东北阔叶红松林群落类型划分及物种多样性.应用生态学报,2007,18(11):2406-2411.
[18]吴晓莆,朱彪,赵淑清,朴世龙,方精云.东北地区阔叶红松林的群落结构及其物种多样性比较.生物多样性,2004,12(1):174-181.
[19]方精云,王襄平,沈泽昊,唐志尧,贺金生,于丹,江源,王志恒,郑成洋,朱江玲,郭兆迪.植物群落清查的主要内容、方法和技术规范.生物多样性,2009,17(6):533-548.
[20]赖江山,米湘成,任海保,马克平.基于多元回归树的常绿阔叶林群丛数量分类:以古田山24公顷森林样地为例.植物生态学报,2010,34(7):761-769.
[21]陈云,王海亮,韩军旺,韦博良,贾宏汝,叶永忠,袁志良.小秦岭森林群落数量分类、排序及多样性垂直格局.生态学报,2014,34(8):2068-2075.
[22] Borcard D,Gillet F,Legendre P.数量生态学——R语言的应用.赖江山,译.北京:高等教育出版社,2014:90-97.
[23] Vittoz P,Bayfield N,Brooker R,Elston D A,Duff E I,Theurillat J P,Guisan A. Reproducibility of species lists,visual cover estimates and frequency methods for recording high-mountain vegetation. Journal of Vegetation Science,2010,21(6):1035-1047.
[24] Zhang J T,Xu B,Li M. Vegetation patterns and species diversity along elevational and disturbance gradients in the Baihua Mountain reserve,Beijing,China. Mountain Research and Development,2013,33(2):170-178.
[25]周以良.中国东北植被地理.北京:科学出版社,1997:16-78.
[26]吴征镒.中国植被.北京:科学出版社,1980:1-1375.
[27]王世雄.黄土高原子午岭植物群落物种多样性的时空格局与过程[D].西安:陕西师范大学,2013.
[28]贺金生,陈伟烈.陆地植物群落物种多样性的梯度变化特征.生态学报,1997,17(1):91-99.
[29]尚文艳,吴钢,付晓,刘阳.陆地植物群落物种多样性维持机制.应用生态学报,2005,16(3):573-578.
[30] Gentry A H. Changes in plant community diversity and floristic composition on environmental and geographical gradients. Annals of the Missouri Botanical Garden,1988,75(1):1-34.
[2]白晓航,张金屯,曹科,王云泉,Sadia S,曹格.河北小五台山国家级自然保护区森林群落与环境的关系.生态学报,2017,37(11):3683-3696.
[3]田永强,俞超超,王磊,黄邦钦.福建九龙江北溪浮游植物群落分布特征及其影响因子.应用生态学报,2012,23(9):2559-2565.
[4]武春华.黄土丘陵区典型植物群落特征及其与环境因子的关系[D].北京:中国科学院研究生院,2008.
[5]赵从举,康慕谊,雷加强.准噶尔盆地典型地段植物群落及其与环境因子的关系.生态学报,2011,31(10):2669-2677.
[6]袁铁象,张合平,欧芷阳,谭一波.地形对桂西南喀斯特山地森林地表植物多样性及分布格局的影响.应用生态学报,2014,25(10):2803-2810.
[7]任萍.黄土高原马栏林区植物群落物种多度研究[D].西安:陕西师范大学,2010.
[8]苏日古嘎,张金屯,张斌,程佳佳,田世广,张钦弟,刘素军.松山自然保护区森林群落的数量分类和排序.生态学报,2010,30(10):2621-2629.
[9]张文静,张钦弟,王晶,冯飞,毕润成.多元回归树与双向指示种分析在群落分类中的应用比较.植物生态学报,2015,39(6):586-592.
[10]张金屯.数量生态学.北京:科学出版社,2011:123-127.
[11] Virtanen R,Luoto M,RmT,Mikkola K,Hjort J,Grytnes J A,Birks H J B. Recent vegetation changes at the high-latitude tree line ecotone are controlled by geomorphological disturbance,productivity and diversity. Global Ecology and Biogeography,2010,19(6):810-821.
[12]李博.生态学.北京:高等教育出版社,2000:168-172.
[13]代力民,谷会岩,邵国凡,王庆礼.中国长白山阔叶红松林.沈阳:辽宁科学技术出版社,2004:5-11.
[14]吴正方,邓慧平.东北阔叶红松林全球气候变化响应研究.地理学报,1996,51(增刊):81-91.
[15]吴正方.东北阔叶红松林分布区生态气候适宜性及全球气候变化影响评价.应用生态学报,2003,14(5):771-775.
[16]贾翔,马芳芳,周旺明,周莉,于大炮,秦静,代力民.气候变化对阔叶红松林潜在地理分布区的影响.生态学报,2017,37(2):464-473.
[17]曹伟,李岩,王树良,于兴华,朱彩霞,刘童燕.东北阔叶红松林群落类型划分及物种多样性.应用生态学报,2007,18(11):2406-2411.
[18]吴晓莆,朱彪,赵淑清,朴世龙,方精云.东北地区阔叶红松林的群落结构及其物种多样性比较.生物多样性,2004,12(1):174-181.
[19]方精云,王襄平,沈泽昊,唐志尧,贺金生,于丹,江源,王志恒,郑成洋,朱江玲,郭兆迪.植物群落清查的主要内容、方法和技术规范.生物多样性,2009,17(6):533-548.
[20]赖江山,米湘成,任海保,马克平.基于多元回归树的常绿阔叶林群丛数量分类:以古田山24公顷森林样地为例.植物生态学报,2010,34(7):761-769.
[21]陈云,王海亮,韩军旺,韦博良,贾宏汝,叶永忠,袁志良.小秦岭森林群落数量分类、排序及多样性垂直格局.生态学报,2014,34(8):2068-2075.
[22] Borcard D,Gillet F,Legendre P.数量生态学——R语言的应用.赖江山,译.北京:高等教育出版社,2014:90-97.
[23] Vittoz P,Bayfield N,Brooker R,Elston D A,Duff E I,Theurillat J P,Guisan A. Reproducibility of species lists,visual cover estimates and frequency methods for recording high-mountain vegetation. Journal of Vegetation Science,2010,21(6):1035-1047.
[24] Zhang J T,Xu B,Li M. Vegetation patterns and species diversity along elevational and disturbance gradients in the Baihua Mountain reserve,Beijing,China. Mountain Research and Development,2013,33(2):170-178.
[25]周以良.中国东北植被地理.北京:科学出版社,1997:16-78.
[26]吴征镒.中国植被.北京:科学出版社,1980:1-1375.
[27]王世雄.黄土高原子午岭植物群落物种多样性的时空格局与过程[D].西安:陕西师范大学,2013.
[28]贺金生,陈伟烈.陆地植物群落物种多样性的梯度变化特征.生态学报,1997,17(1):91-99.
[29]尚文艳,吴钢,付晓,刘阳.陆地植物群落物种多样性维持机制.应用生态学报,2005,16(3):573-578.
[30] Gentry A H. Changes in plant community diversity and floristic composition on environmental and geographical gradients. Annals of the Missouri Botanical Garden,1988,75(1):1-34.