广东康禾自然保护区地面苔藓种群分布与格局分析
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摘要
种群分布格局是植物群落的重要特征,研究种群分布格局是理解植物群落形成和构建的基础。采用Poisson数学模型和方差均值法检验广东康禾自然保护区地面苔藓种群分布格局的类型,利用双向轨迹方差法(TTLQV)分析其格局变化。结果显示,研究区苔藓植物整体呈集群分布,优势种东亚拟鳞叶藓(Pseudotaxiphyllum pohliaecarpum)平均拥挤指数最高(m*=13.04),聚块性指数最低(m*/m=1.15),细叶小羽藓(Haplocladium microphyllum)和细指苔(Kurzia gonyotricha)的平均拥挤指数次之,分别为11.85、10.00;而非优势物种树生扁萼苔(Radula obscura)和长叶羽苔(Plagiochila flexuosa)聚快性指数最高,分别为14.79、14.45;说明优势物种的种间竞争更强烈,非优势物种分布更聚集。种群格局变化表明,研究区苔藓植物的格局规模以小斑块聚集居多,如细指苔、神山细鳞苔(Lejeunea eifrigii)、东亚拟鳞叶藓及南方小锦藓(Brotherella henonii)等多个物种在区组4、5、15、22、24呈现斑块聚集,其格局规模的差异可能与苔藓植物的生存策略和生长环境密切相关。苔类植物与苔藓植物的整体格局变化更接近,在区组4都呈大斑块聚集,整体为镶嵌格局;而藓类植物在区组5、16、22均有较大的峰值波动,为大斑块聚集;说明苔类植物优势种群组在群落中占主导地位。不同苔藓物种呈现不同的格局变化,主要受其生物学特性与种间竞争的影响,生境异质性则是其集群分布的主要原因。
Population distribution patterns are the key characteristics of plant communities. Investigating population distribution patterns is fundamental to the understanding of the formation and assembly of plant communities. Here, we conducted a study on the population distribution patterns of ground bryophytes in Kanghe Nature Reserve of Guangdong Province, and tested various patterns using Poisson Model and the variance/mean approach. We also analyzed their pattern changes with Two-term local quadrat variance(TTLQV) technique. The results showed that most bryophyte species were distributed in a clumped pattern, and the dominant species Pseudotaxiphyllum pohliaecarpum had the highest mean crowding index(m*=13.04) and the lowest clustering index(m*/m=1.15), followed by Haplocladium microphyllum and Kurzia gonyotricha with a mean crowding index of 11.85 and 10, respectively; while the two non-dominant species, Radula obscura and Plagiochila flexuosa, had the highest patchiness index of 14.79 and 14.45, respectively. These results suggested that the dominant species had stronger interspecific competition than non-dominant species, while the non-dominant species followed a more aggregated distribution pattern. Changes in population distribution patterns indicated that the pattern scale of the bryophytes was dominated by small sizes in the study area. Several species were clumped at block 4, 5, 15, 22, and 24, such as Kurzia gonyotricha, Lejeunea eifrigii, Pseudotaxiphyllum pohliaecarpum, Brotherella henonii. The scale difference in their distribution patterns was closely related to the survival strategies and habitats of the bryophytes. The pattern changes of liverworts and bryophytes were more similar at block 4, with large patches and an overall mosaic pattern; while the mosses had large patches at block 5, 16 and 22, due to larger peak fluctuations. These results indicate that the dominant populations of liverworts play leading role in the community assembly. Different bryophytes exhibite different pattern changes, which are mainly affected by their biological characteristics and interspecific competition, while habitat heterogeneity is chiefly responsible for the clumped distribution of bryophytes.
Population distribution patterns are the key characteristics of plant communities. Investigating population distribution patterns is fundamental to the understanding of the formation and assembly of plant communities. Here, we conducted a study on the population distribution patterns of ground bryophytes in Kanghe Nature Reserve of Guangdong Province, and tested various patterns using Poisson Model and the variance/mean approach. We also analyzed their pattern changes with Two-term local quadrat variance(TTLQV) technique. The results showed that most bryophyte species were distributed in a clumped pattern, and the dominant species Pseudotaxiphyllum pohliaecarpum had the highest mean crowding index(m*=13.04) and the lowest clustering index(m*/m=1.15), followed by Haplocladium microphyllum and Kurzia gonyotricha with a mean crowding index of 11.85 and 10, respectively; while the two non-dominant species, Radula obscura and Plagiochila flexuosa, had the highest patchiness index of 14.79 and 14.45, respectively. These results suggested that the dominant species had stronger interspecific competition than non-dominant species, while the non-dominant species followed a more aggregated distribution pattern. Changes in population distribution patterns indicated that the pattern scale of the bryophytes was dominated by small sizes in the study area. Several species were clumped at block 4, 5, 15, 22, and 24, such as Kurzia gonyotricha, Lejeunea eifrigii, Pseudotaxiphyllum pohliaecarpum, Brotherella henonii. The scale difference in their distribution patterns was closely related to the survival strategies and habitats of the bryophytes. The pattern changes of liverworts and bryophytes were more similar at block 4, with large patches and an overall mosaic pattern; while the mosses had large patches at block 5, 16 and 22, due to larger peak fluctuations. These results indicate that the dominant populations of liverworts play leading role in the community assembly. Different bryophytes exhibite different pattern changes, which are mainly affected by their biological characteristics and interspecific competition, while habitat heterogeneity is chiefly responsible for the clumped distribution of bryophytes.
引文
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DAI X,EVANDER M.1997.Transect-based patch size frequency analysis[J].Journal of Vegetation Science,8(6):865-872.
EBERT A,DA COSTA R B,BRONDANI G E.2016.Spatial distribution pattern of Mezilaurus itauba(Meins.)Taub.Ex mez.in a seasonal forest area of the southern Amazon,Brazil[J].Iforest-Biogeosciences and Forestry,9:497-502.
FIBICH P,LEPS J,NOVOTNY V,et al.2016.Spatial patterns of tree species distribution in New Guinea primary and secondary lowland rain forest[J].Journal of Vegetation Science,27(2):328-339.
HANEWINKEL M.2004.Spatial patterns in mixed coniferous even-aged,uneven-aged and conversion stands[J].European Journal of Forest Research,123(2):139-155.
HE S Y,ZHONG Y L,SUN Y D,et al.2017.Topography-associated thermal gradient predicts warming effects on woody plant structural diversity in a subtropical forest[J].Scientific Reports,7:40387.
HE X L,HE K S,HYVONEN J.2016.Will bryophytes survive in a warming world[J].Perspectives in Plant Ecology Evolution and Systematics,19:49-60.
KREBS C.1989.Ecological Methodology[M].New York:Harper Collins publishers.
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MILLER H A.1982.Bryophyte evolution and geography[J].Biological Journal of the Linnean Society,18:145-196.
NGUYEN H H,URIA-DIEZ J,WIEGAND K.2016.Spatial distribution and association patterns in a tropical evergreen broad-leaved forest of north-central Vietnam[J].Journal of Vegetation Science,27(2):318-327.
BATISTA W V S M,P?RTO K C,SANTOS N D D.2018.Distribution,ecology,and reproduction of bryophytes in a humid enclave in the semiarid region of northeastern Brazil[J].Acta Botanica Brasilica,32(2):303-313.
SONG L,LU H Z,XU X L,et al.2016.Organic nitrogen uptake is a significant contributor to nitrogen economy of subtropical epiphytic bryophytes[J].Scientific Reports,6:30408.
SONG L,MA W Z,YAO Y L,et al.2015.Bole bryophyte diversity and distribution patterns along three altitudinal gradients in Yunnan,China[J].Journal of Vegetation Science,26(3):576-587.
SZMYT J.2014.Spatial statistics in ecological analysis:from indices to functions[J].Silva Fennica,48(1):93-109.
USHER M B.1983.Pattern in the Simple Moss-Turf Communities of the Sub-Antarctic and Maritime Antarctic[J].Journal of Ecology,71(3):945-958.
YANG F,ZHOU Y.2017.Quantifying spatial scale of positive and negative terrains pattern at watershed-scale:Case in soil and water conservation region on Loess Plateau[J].Journal of Mountain Science,14(8):1642-1654.
艾应伟,刘浩,李伟,等.2010.苔藓植物资源保护利用中的生态环境效应研究进展[J].生态环境学报,19(1):227-232.
卜兆君,陈旭,姜丽红,等.2009.苔藓植物相互作用的研究进展[J].应用生态学报,20(2):460-466.
陈勤,孙冲,方炎明.2013.苔藓植物的生态环境指示作用[J].世界林业研究,26(2):19-23.
陈云,冯佳伟,牛帅,等.2017.小秦岭自然保护区苔藓植物群落数量分类、排序及多样性垂直格局[J].生态学报,37(8):2653-2664.
董秋莲,李毅,单立山,等.2013.张掖市龙首山红砂种群结构和分布格局研究[J].水土保持通报,33(4):284-288.
郭磊,韦博良,胡金涛,等.2016.基于两个不同资源轴上苔藓植物生态位分析[J].生态学报,21(37):7266-7276.
郭连金.2014.濒危植物香果树幼苗空间格局及数量动态研究[J].西北植物学报,34(9):1887-1893.
贾成朕,苗百岭,姜威,等.2017.阿拉善典型荒漠群落种群空间格局及种间关联的尺度依赖性[J].内蒙古大学学报(自然版),48(2):142-148.
贾渝,何思.2013.中国生物物种名录:苔藓植物[M].北京:科学出版社.
姜苹红,罗远玲,彭克俭,等.2015.苔藓植物运用于大气重金属污染监测的研究进展[J].环境污染与防治,37(7):82-87.
兰国玉,雷瑞德.2003.植物种群空间分布格局研究方法概述[J].西北林学院学报,18(2):17-21.
李海涛.1995.植物种群分布格局研究概况[J].植物学报,12(2):19-26.
李素清,李斌,张金屯.2006.基于DCA-TTLQV的云顶山亚高山草甸群落格局分析[J].草业学报,15(6):44-48.
李素清,武冬梅,王涛,等.2011.山西长治湿地草本植物优势种群和群落的空间格局分析[J].草业学报,20(3):43-50.
李亚,胡小柯,魏怀东,等.2017.腾格里沙漠南缘天然群落主要种群空间分布格局研究[J].西北林学院学报,32(2):67-72.
刘蔚秋,雷纯义,戴小华.2007.广东黑石顶森林苔藓植物群落特征初探[J].热带亚热带植物学报,15(6):538-544.
马钦彦.2009.负二项式分布及种群格局检验分析[J].北京林业大学学报,31(3):1-5.
梅军林,庄枫红,马姜明,等.2017.桂林喀斯特地区克隆生长红背山麻杆种群的点格局分析[J].生态学报,37(9):3164-3171.
田维莉,孙守琴.2011.苔藓植物生态功能研究新进展[J].生态学杂志,30(6):1265-1269.
王继和,靳虎甲,马全林,等.2010.干旱区油蒿种群结构和分布格局分析[J].中国沙漠,30(3):534-538.
王利松,贾渝,张宪春,等.2015.中国高等植物多样性[J].生物多样性,23(2):217-224.
王中生,安树青,方炎明.2003.苔藓植物生殖生态学研究[J].生态学报,23(11):2444-2452.
吴德邻.2013.广东苔藓志[M].广州:广东科技出版社.
许宁,韦博良,胡金涛,等.2017.宝天曼自然保护区树附生苔藓空间格局分析[J].河南农业大学学报,51(2):230-236.
张金屯.2005.芦芽山亚高山草甸优势种群和群落的二维格局分析[J].生态学报,25(6):1264-1268.
张金屯.2011.数量生态学[M].第2版.北京:科学出版社.
张金屯,孟东平.2006.芦芽山油松-辽东栎林优势树种空间分布格局研究[J].西北植物学报,26(8):1682-1685.
张瑾,陈文业,张继强,等.2013.甘肃敦煌西湖荒漠-湿地生态系统优势植物种群分布格局及种间关联性[J].中国沙漠,33(2):349-357.
张维,李海燕,赖晓辉,等.2016.新疆天山峡谷不同坡向野核桃种群分布格局[J].应用生态学报,27(10):3105-3113.
张旭,李家湘,喻勋林,等.2015.湖南大围山杜鹃灌丛木本植物种群空间格局[J].生态学杂志,34(11):3034-3039.
张元明,曹同,潘伯荣.2002.干旱与半干旱地区苔藓植物生态学研究综述[J].生态学报,22(7):1129-1134.
赵峰侠,尹林克.2007.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[J].生态学杂志,26(7):972-977.
DA COSTA D P,DOS SANTOS N D,DE REZENDE M A,et al.2015.Bryoflora of the Itatiaia National Park along an elevation gradient:diversity and conservation[J].Biodiversity and Conservation,24(9):2199-2212.
DAI X,EVANDER M.1997.Transect-based patch size frequency analysis[J].Journal of Vegetation Science,8(6):865-872.
EBERT A,DA COSTA R B,BRONDANI G E.2016.Spatial distribution pattern of Mezilaurus itauba(Meins.)Taub.Ex mez.in a seasonal forest area of the southern Amazon,Brazil[J].Iforest-Biogeosciences and Forestry,9:497-502.
FIBICH P,LEPS J,NOVOTNY V,et al.2016.Spatial patterns of tree species distribution in New Guinea primary and secondary lowland rain forest[J].Journal of Vegetation Science,27(2):328-339.
HANEWINKEL M.2004.Spatial patterns in mixed coniferous even-aged,uneven-aged and conversion stands[J].European Journal of Forest Research,123(2):139-155.
HE S Y,ZHONG Y L,SUN Y D,et al.2017.Topography-associated thermal gradient predicts warming effects on woody plant structural diversity in a subtropical forest[J].Scientific Reports,7:40387.
HE X L,HE K S,HYVONEN J.2016.Will bryophytes survive in a warming world[J].Perspectives in Plant Ecology Evolution and Systematics,19:49-60.
KREBS C.1989.Ecological Methodology[M].New York:Harper Collins publishers.
MANDL N A,KESSLER M,GRADSTEIN S R.2009.Effects of environmental heterogeneity on species diversity and composition of terrestrial bryophyte assemblages in tropical montane forests of southern Ecuador[J].Plant Ecology&Diversity,2(3):313-321.
MILLER H A.1982.Bryophyte evolution and geography[J].Biological Journal of the Linnean Society,18:145-196.
NGUYEN H H,URIA-DIEZ J,WIEGAND K.2016.Spatial distribution and association patterns in a tropical evergreen broad-leaved forest of north-central Vietnam[J].Journal of Vegetation Science,27(2):318-327.
BATISTA W V S M,P?RTO K C,SANTOS N D D.2018.Distribution,ecology,and reproduction of bryophytes in a humid enclave in the semiarid region of northeastern Brazil[J].Acta Botanica Brasilica,32(2):303-313.
SONG L,LU H Z,XU X L,et al.2016.Organic nitrogen uptake is a significant contributor to nitrogen economy of subtropical epiphytic bryophytes[J].Scientific Reports,6:30408.
SONG L,MA W Z,YAO Y L,et al.2015.Bole bryophyte diversity and distribution patterns along three altitudinal gradients in Yunnan,China[J].Journal of Vegetation Science,26(3):576-587.
SZMYT J.2014.Spatial statistics in ecological analysis:from indices to functions[J].Silva Fennica,48(1):93-109.
USHER M B.1983.Pattern in the Simple Moss-Turf Communities of the Sub-Antarctic and Maritime Antarctic[J].Journal of Ecology,71(3):945-958.
YANG F,ZHOU Y.2017.Quantifying spatial scale of positive and negative terrains pattern at watershed-scale:Case in soil and water conservation region on Loess Plateau[J].Journal of Mountain Science,14(8):1642-1654.
艾应伟,刘浩,李伟,等.2010.苔藓植物资源保护利用中的生态环境效应研究进展[J].生态环境学报,19(1):227-232.
卜兆君,陈旭,姜丽红,等.2009.苔藓植物相互作用的研究进展[J].应用生态学报,20(2):460-466.
陈勤,孙冲,方炎明.2013.苔藓植物的生态环境指示作用[J].世界林业研究,26(2):19-23.
陈云,冯佳伟,牛帅,等.2017.小秦岭自然保护区苔藓植物群落数量分类、排序及多样性垂直格局[J].生态学报,37(8):2653-2664.
董秋莲,李毅,单立山,等.2013.张掖市龙首山红砂种群结构和分布格局研究[J].水土保持通报,33(4):284-288.
郭磊,韦博良,胡金涛,等.2016.基于两个不同资源轴上苔藓植物生态位分析[J].生态学报,21(37):7266-7276.
郭连金.2014.濒危植物香果树幼苗空间格局及数量动态研究[J].西北植物学报,34(9):1887-1893.
贾成朕,苗百岭,姜威,等.2017.阿拉善典型荒漠群落种群空间格局及种间关联的尺度依赖性[J].内蒙古大学学报(自然版),48(2):142-148.
贾渝,何思.2013.中国生物物种名录:苔藓植物[M].北京:科学出版社.
姜苹红,罗远玲,彭克俭,等.2015.苔藓植物运用于大气重金属污染监测的研究进展[J].环境污染与防治,37(7):82-87.
兰国玉,雷瑞德.2003.植物种群空间分布格局研究方法概述[J].西北林学院学报,18(2):17-21.
李海涛.1995.植物种群分布格局研究概况[J].植物学报,12(2):19-26.
李素清,李斌,张金屯.2006.基于DCA-TTLQV的云顶山亚高山草甸群落格局分析[J].草业学报,15(6):44-48.
李素清,武冬梅,王涛,等.2011.山西长治湿地草本植物优势种群和群落的空间格局分析[J].草业学报,20(3):43-50.
李亚,胡小柯,魏怀东,等.2017.腾格里沙漠南缘天然群落主要种群空间分布格局研究[J].西北林学院学报,32(2):67-72.
刘蔚秋,雷纯义,戴小华.2007.广东黑石顶森林苔藓植物群落特征初探[J].热带亚热带植物学报,15(6):538-544.
马钦彦.2009.负二项式分布及种群格局检验分析[J].北京林业大学学报,31(3):1-5.
梅军林,庄枫红,马姜明,等.2017.桂林喀斯特地区克隆生长红背山麻杆种群的点格局分析[J].生态学报,37(9):3164-3171.
田维莉,孙守琴.2011.苔藓植物生态功能研究新进展[J].生态学杂志,30(6):1265-1269.
王继和,靳虎甲,马全林,等.2010.干旱区油蒿种群结构和分布格局分析[J].中国沙漠,30(3):534-538.
王利松,贾渝,张宪春,等.2015.中国高等植物多样性[J].生物多样性,23(2):217-224.
王中生,安树青,方炎明.2003.苔藓植物生殖生态学研究[J].生态学报,23(11):2444-2452.
吴德邻.2013.广东苔藓志[M].广州:广东科技出版社.
许宁,韦博良,胡金涛,等.2017.宝天曼自然保护区树附生苔藓空间格局分析[J].河南农业大学学报,51(2):230-236.
张金屯.2005.芦芽山亚高山草甸优势种群和群落的二维格局分析[J].生态学报,25(6):1264-1268.
张金屯.2011.数量生态学[M].第2版.北京:科学出版社.
张金屯,孟东平.2006.芦芽山油松-辽东栎林优势树种空间分布格局研究[J].西北植物学报,26(8):1682-1685.
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