天坑洞穴群苔类群落分布规律与环境因子关系——以贵州猴耳天坑为例
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摘要
了解天坑内洞穴苔类群落分布规律与环境因子的关系,是天坑内洞穴生物多样性保护的重要步骤。本研究以贵州猴耳天坑不同深度4个洞穴苔类群落为对象,进行了苔类数据的多元回归分析、α和β多样性分析、苔类和环境因子典范对应分析,结果表明:猴耳天坑洞穴内苔类群落组成单一,多为单优势种或双优势种群落。垂直方向上,猴耳天坑洞穴苔类群落物种数随天坑深度加深呈增加趋势。经交叉验证,天坑内4个洞穴苔类群落呈现梯度性分布,可分成三类,Ⅰ:天坑中部洞穴群落;主要为艳绿光苔(Cyathodium smaragdi-num)+花叶溪苔(Pellia endiviifolia)+叉钱苔(Riccia fluitans)群落。Ⅱ:天坑中下部洞穴群落;主要为艳绿光苔(Cyathodium smaragdinum)+叉钱苔(Riccia fluitans)群落。Ⅲ:天坑底部洞穴群落;主要为花叶溪苔(Pellia endiviifolia)+小蛇苔(Conocephalum japonicum)+毛地钱(Dumortiera hirsuta)群落。水平方向上,万亩梯田洞穴苔类群落物种多样性变化与其群落物种组成先增加后减小的变化趋势保持一致,群落环境异质性呈现先减小后增大的趋势;天乐堂洞穴苔类群落物种多样性和其环境异质性变化保持一致,群落物种多样性先减小后增加,群落环境异质性先减小后增大。典范对应分析显示,苔类群落在垂直和水平方向的分布受到温度、湿度和光照的梯度性影响,3个环境因子的不同组合影响苔类群落的多样性(第一轴解释量45.89%,第二轴79.28%,P<0.05)。
Revealing the relationship between distribution pattern of liverworts community and environmental factors is an important step for biodiversity conservation in Tiankeng cave. Multivariate regression trees analysis,α-and β-diversity analysis,and canonical correspondence analysis were conducted to analyze the distribution patterns of liverwort communities in four caves with different depths in Monkey-Ear Tiankeng,Guizhou Province and their relationships with environmental factors. The results showed that the community composition of liverworts was simple in Tiankeng caves,mainly consisting of one or two dominant species. Species richness of liverwort community increased with the increasing depth of cave location,showing a gradient pattern. The liverwort communities could be divided into three types from four caves using cross-validation:( Ⅰ) Communities of middle-depth cave,mainly consisting of Cyathodium smaragdinum+Pellia endiviifolia+Riccia fluitans;( Ⅱ) Communities of lower-middle-depth cave,mainly consisting of Cyathodium smaragdinum+Riccia fluitans;( Ⅲ) Communities of bottom cave,mainly including Pellia endiviifolia + Conocephalum japonicum + Dumortiera hirsuta. In horizontal direction,the diversity of liverwort community changed with community composition( species,genera,and families) in the Wanmutitian cave,which was first increased and then declined. However,the environmental heterogeneity of communities showed a contrasting pattern. The diversity of liverworts was consistent with the environmental heterogeneity in the Tianletang cave,which was first decreased and then increased. The distributions of liverwort community in both vertical and horizontal directions were affected by the gradient variations of temperature,humidity,and illumination. The diversity of liverwort community was a result of the combinations of three environmental factors. The interpretation of axis 1 and axis 2 was 45. 89% and 79. 28% from canonical correspondence analysis,respectively.
Revealing the relationship between distribution pattern of liverworts community and environmental factors is an important step for biodiversity conservation in Tiankeng cave. Multivariate regression trees analysis,α-and β-diversity analysis,and canonical correspondence analysis were conducted to analyze the distribution patterns of liverwort communities in four caves with different depths in Monkey-Ear Tiankeng,Guizhou Province and their relationships with environmental factors. The results showed that the community composition of liverworts was simple in Tiankeng caves,mainly consisting of one or two dominant species. Species richness of liverwort community increased with the increasing depth of cave location,showing a gradient pattern. The liverwort communities could be divided into three types from four caves using cross-validation:( Ⅰ) Communities of middle-depth cave,mainly consisting of Cyathodium smaragdinum+Pellia endiviifolia+Riccia fluitans;( Ⅱ) Communities of lower-middle-depth cave,mainly consisting of Cyathodium smaragdinum+Riccia fluitans;( Ⅲ) Communities of bottom cave,mainly including Pellia endiviifolia + Conocephalum japonicum + Dumortiera hirsuta. In horizontal direction,the diversity of liverwort community changed with community composition( species,genera,and families) in the Wanmutitian cave,which was first increased and then declined. However,the environmental heterogeneity of communities showed a contrasting pattern. The diversity of liverworts was consistent with the environmental heterogeneity in the Tianletang cave,which was first decreased and then increased. The distributions of liverwort community in both vertical and horizontal directions were affected by the gradient variations of temperature,humidity,and illumination. The diversity of liverwort community was a result of the combinations of three environmental factors. The interpretation of axis 1 and axis 2 was 45. 89% and 79. 28% from canonical correspondence analysis,respectively.
引文
陈邦杰.1958.中国苔藓植物生态群落和地理分布初步报告.植物分类学报,7(4):270-293.
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.生态学报,37(8):2653-2664.
戈峰.2008.现代生态学.北京:科学出版社.
高谦,吴玉环.2010.中国苔纲和角苔纲植物属志.北京:科学出版社.
黄文琥,王之明,梁盛.2014.不同生境苔类和角苔类植物对生态环境的指示作用.中国环境监测,(3):70-74.
姬明飞,韩鸿基.2017.两种藓类植物对光强的适应性差异.草业科学,34(9):1787-1792.
简小枚,税伟,王亚楠,等.2018.重度退化的喀斯特天坑草地物种多样性及稳定性研究---以云南沾益退化天坑为例.生态学报,38(13):4704-4714.
赖江山,米湘成,任海保,等.2010.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,34(7):761-769.
矢佳昱,韩海荣,程小琴,等.2018.辽河源自然保护区不同年龄天然油松林林下草本层植物物种多样性及其环境解释.生态学杂志,37(5):1326-1333.
税伟,陈毅萍,简小枚,等.2018.喀斯特原生天坑垂直梯度上植物多样性特征---以云南沾益天坑为例.山地学报,36(1):53-62.
苏宇乔,薛跃规,范蓓蓓,等.2016.广西流星天坑植物群落结构与多样性.西北植物学报,36(11):2300-2306.
唐礼俊,杨思河,林继惠,等.1998.长白山藓类植物在失水干燥及再水化过程中的CO2同化能力和呼吸速率的变化.应用与环境生物学报,4(1):6-9.
唐启明,薛跃规,党桂兰,等.2016.广西苔藓植物新纪录.广西师范大学学报:自然科学版,34(2):143-146.
张朝晖,钟本固,王智慧,等.1996.黄果树喀斯特洞穴群苔藓植物群落.贵州科学,14(1):37-46.
张朝晖,祝安.1996b.黄果树喀斯特洞穴群苔藓植物岩溶的初步研究.中国岩溶,15(3):224-232.
张朝晖,艾伦·培特客斯.2002.英格兰洞穴苔藓植物区系特征及其岩溶沉积研究.西北植物学报,22(2):359-367.
张朝晖,赵传海,李晓娜,等.2005.中国桂林岩溶洞穴苔藓植物研究.广西植物,25(2):107-111.
Ah-Peng C,Cardoso AW,Flores O,et al.2017.The role of epiphytic bryophytes in interception,storage,and the regulated release of atmospheric moisture in a tropical montane cloud forest.Journal of Hydrology,548:665-673.
Ammons N.1933.Bryophytes of McKinney’s cave.Bryologist,36:16-19.
Ariyanti NS,Bos MM,Kartawinata K,et al.2008.Bryophytes on tree trunks in natural forests,selectively logged forests and cacao agroforests in central Sulawesi,Indonesia.Biological Conservation,141:2516-2527.
Burney DA,Burney LP.2016.Monitoring results from a decade of native plant translocations at Makauwahi Cave Reserve,Kaua‘i.Plant Ecology,217:139-153.
Da SS,Mrquez JRG,Sommer JH,et al.2018.Plant biodiversity patterns along a climatic gradient and across protected areas in West Africa.African Journal of Ecology,56:641-652.
Field KJ,Rimington WR,Bidartondo MI,et al.2016.Functional analysis of liverworts in dual symbiosis with Glomeromycota and Mucoromycotina fungi under a simulated Palaeozoic CO2decline.ISME Journal,10:1514-1526.
Fraser CI,Connell L,Lee CK,et al.2018.Evidence of plant and animal communities at exposed and subglacial(cave)geothermal sites in Antarctica.Polar Biology,41:417-421.
Hattaway RA.1980.The calciphilous bryophytes of three limestone sinks in eastern Tennessee.Bryologist,83:161-169.
Janik E,Bednarska J,Zubik M,et al.2017.A chloroplast“wake up”mechanism:Illumination with weak light activates the photosynthetic antenna function in dark-adapted plants.Journal of Plant Physiology,210:1-8.
KuglerovL,Dynesius M,Laudon H,et al.2016.Relationships between plant assemblages and water flow across a boreal forest landscape:A comparison of liverworts,mosses,and vascular plants.Ecosystems,19:170-184.
Li J,Liu Y,Cao T,et al.1999.A comparative study on ecophysiological characteristics of the two overwintering host mosses of gallaphids.Nova Hedwigia,68:233-240.
Longton RE.1979.Vegetation ecology and classification in the maritime Antarctic zone.Canadian Journal of Botany,57:2264-2278.
Lbel S,Mair L,Lnnell N,et al.2018.Biological traits explain bryophyte species distributions and responses to forest fragmentation and climatic variation.Journal of Ecology,106:1700-1713.
Magurran AE.1989.Ecological diversity and its measurement.Biometrics,46:81-99.
Mason-Williams M,Benson-Evans K.1967.Summary of results obtained during a preliminary investigation into the bacterial and botanical flora of caves in south Wales.International Journal of Speleology,2:397-402.
Miller NG,Vitt DH.1970.Additional bryophytes from sinkholes in Alpena country,Michigan,including Orthotrichum pallens new to eastern North America.Michigan Botanist,2:369-376.
Morris JL,Puttick MN,Clark JW,et al.2018.The timescale of early land plant evolution.Proceedings of the National Academy of Sciences of the United States of America,115:E2274-E2283.
Pentecost A,Zhang ZH.2001.The distribution of plants in Scoska cave,north Yorkshire,and their relationship to light intensity.International Journal of Speleology,30:2-7.
Reyes-Colón CM,Sastre DJI.2000.Bryophytes of two sinkholes of the north-central karst zone of Puerto Rico.Evansia,17:68-76.
Rincón E.1993.Growth responses of six bryophyte species to different light intensities.Canadian Journal of Botany,71:661-665.
Robinson H,Wells J.1956.The bryophytes of certain limestone sinks in Alpena County,Michigan.Bryologist,59:12-17.
Schluter D,Pennell MW.2017.Speciation gradients and the distribution of biodiversity.Nature,546:48-55.
Shen L,Guo SL,Song HT,et al.2011.Influence of sample size on bryophyte ecological indices.Guihaia,31:198-203.
Smilauer P,LepJ.2005.Multivariate analysis of ecological data using CANOCO.Bulletin of the Ecological Society of America,86:201-201.
Stehn SE,Webster CR,Glime JM,et al.2010.Elevational gradients of bryophyte diversity,life forms,and community assemblage in the southern Appalachian Mountains.Canadian Journal of Forest Research,40:2164-2174.
Thatcher EP.1949.Bryophytes of an artificially illuminated cave.Bryologist,52:212-214.
Woziwoda B,Staniaszek-Kik M,Stefańska-Krzaczek E.2017.Assemblages of native bryophytes in secondary forests with introduced Quercus rubra.Nordic Journal of Botany,35:1-10.
Zhang ZH,Peng T,Li XN,et al.2004.A study on the bryophytes of karst cave thresh-hold at Kunming area in Yunnan province,P.R.China.Carsologica Sinica,23:229-234.
Zhu XW,Tony W.2006.Tiankeng:Definition and description.Speleogenesis&Evolution of Karst Aquifers,4:8.
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.生态学报,37(8):2653-2664.
戈峰.2008.现代生态学.北京:科学出版社.
高谦,吴玉环.2010.中国苔纲和角苔纲植物属志.北京:科学出版社.
黄文琥,王之明,梁盛.2014.不同生境苔类和角苔类植物对生态环境的指示作用.中国环境监测,(3):70-74.
姬明飞,韩鸿基.2017.两种藓类植物对光强的适应性差异.草业科学,34(9):1787-1792.
简小枚,税伟,王亚楠,等.2018.重度退化的喀斯特天坑草地物种多样性及稳定性研究---以云南沾益退化天坑为例.生态学报,38(13):4704-4714.
赖江山,米湘成,任海保,等.2010.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,34(7):761-769.
矢佳昱,韩海荣,程小琴,等.2018.辽河源自然保护区不同年龄天然油松林林下草本层植物物种多样性及其环境解释.生态学杂志,37(5):1326-1333.
税伟,陈毅萍,简小枚,等.2018.喀斯特原生天坑垂直梯度上植物多样性特征---以云南沾益天坑为例.山地学报,36(1):53-62.
苏宇乔,薛跃规,范蓓蓓,等.2016.广西流星天坑植物群落结构与多样性.西北植物学报,36(11):2300-2306.
唐礼俊,杨思河,林继惠,等.1998.长白山藓类植物在失水干燥及再水化过程中的CO2同化能力和呼吸速率的变化.应用与环境生物学报,4(1):6-9.
唐启明,薛跃规,党桂兰,等.2016.广西苔藓植物新纪录.广西师范大学学报:自然科学版,34(2):143-146.
张朝晖,钟本固,王智慧,等.1996.黄果树喀斯特洞穴群苔藓植物群落.贵州科学,14(1):37-46.
张朝晖,祝安.1996b.黄果树喀斯特洞穴群苔藓植物岩溶的初步研究.中国岩溶,15(3):224-232.
张朝晖,艾伦·培特客斯.2002.英格兰洞穴苔藓植物区系特征及其岩溶沉积研究.西北植物学报,22(2):359-367.
张朝晖,赵传海,李晓娜,等.2005.中国桂林岩溶洞穴苔藓植物研究.广西植物,25(2):107-111.
Ah-Peng C,Cardoso AW,Flores O,et al.2017.The role of epiphytic bryophytes in interception,storage,and the regulated release of atmospheric moisture in a tropical montane cloud forest.Journal of Hydrology,548:665-673.
Ammons N.1933.Bryophytes of McKinney’s cave.Bryologist,36:16-19.
Ariyanti NS,Bos MM,Kartawinata K,et al.2008.Bryophytes on tree trunks in natural forests,selectively logged forests and cacao agroforests in central Sulawesi,Indonesia.Biological Conservation,141:2516-2527.
Burney DA,Burney LP.2016.Monitoring results from a decade of native plant translocations at Makauwahi Cave Reserve,Kaua‘i.Plant Ecology,217:139-153.
Da SS,Mrquez JRG,Sommer JH,et al.2018.Plant biodiversity patterns along a climatic gradient and across protected areas in West Africa.African Journal of Ecology,56:641-652.
Field KJ,Rimington WR,Bidartondo MI,et al.2016.Functional analysis of liverworts in dual symbiosis with Glomeromycota and Mucoromycotina fungi under a simulated Palaeozoic CO2decline.ISME Journal,10:1514-1526.
Fraser CI,Connell L,Lee CK,et al.2018.Evidence of plant and animal communities at exposed and subglacial(cave)geothermal sites in Antarctica.Polar Biology,41:417-421.
Hattaway RA.1980.The calciphilous bryophytes of three limestone sinks in eastern Tennessee.Bryologist,83:161-169.
Janik E,Bednarska J,Zubik M,et al.2017.A chloroplast“wake up”mechanism:Illumination with weak light activates the photosynthetic antenna function in dark-adapted plants.Journal of Plant Physiology,210:1-8.
KuglerovL,Dynesius M,Laudon H,et al.2016.Relationships between plant assemblages and water flow across a boreal forest landscape:A comparison of liverworts,mosses,and vascular plants.Ecosystems,19:170-184.
Li J,Liu Y,Cao T,et al.1999.A comparative study on ecophysiological characteristics of the two overwintering host mosses of gallaphids.Nova Hedwigia,68:233-240.
Longton RE.1979.Vegetation ecology and classification in the maritime Antarctic zone.Canadian Journal of Botany,57:2264-2278.
Lbel S,Mair L,Lnnell N,et al.2018.Biological traits explain bryophyte species distributions and responses to forest fragmentation and climatic variation.Journal of Ecology,106:1700-1713.
Magurran AE.1989.Ecological diversity and its measurement.Biometrics,46:81-99.
Mason-Williams M,Benson-Evans K.1967.Summary of results obtained during a preliminary investigation into the bacterial and botanical flora of caves in south Wales.International Journal of Speleology,2:397-402.
Miller NG,Vitt DH.1970.Additional bryophytes from sinkholes in Alpena country,Michigan,including Orthotrichum pallens new to eastern North America.Michigan Botanist,2:369-376.
Morris JL,Puttick MN,Clark JW,et al.2018.The timescale of early land plant evolution.Proceedings of the National Academy of Sciences of the United States of America,115:E2274-E2283.
Pentecost A,Zhang ZH.2001.The distribution of plants in Scoska cave,north Yorkshire,and their relationship to light intensity.International Journal of Speleology,30:2-7.
Reyes-Colón CM,Sastre DJI.2000.Bryophytes of two sinkholes of the north-central karst zone of Puerto Rico.Evansia,17:68-76.
Rincón E.1993.Growth responses of six bryophyte species to different light intensities.Canadian Journal of Botany,71:661-665.
Robinson H,Wells J.1956.The bryophytes of certain limestone sinks in Alpena County,Michigan.Bryologist,59:12-17.
Schluter D,Pennell MW.2017.Speciation gradients and the distribution of biodiversity.Nature,546:48-55.
Shen L,Guo SL,Song HT,et al.2011.Influence of sample size on bryophyte ecological indices.Guihaia,31:198-203.
Smilauer P,LepJ.2005.Multivariate analysis of ecological data using CANOCO.Bulletin of the Ecological Society of America,86:201-201.
Stehn SE,Webster CR,Glime JM,et al.2010.Elevational gradients of bryophyte diversity,life forms,and community assemblage in the southern Appalachian Mountains.Canadian Journal of Forest Research,40:2164-2174.
Thatcher EP.1949.Bryophytes of an artificially illuminated cave.Bryologist,52:212-214.
Woziwoda B,Staniaszek-Kik M,Stefańska-Krzaczek E.2017.Assemblages of native bryophytes in secondary forests with introduced Quercus rubra.Nordic Journal of Botany,35:1-10.
Zhang ZH,Peng T,Li XN,et al.2004.A study on the bryophytes of karst cave thresh-hold at Kunming area in Yunnan province,P.R.China.Carsologica Sinica,23:229-234.
Zhu XW,Tony W.2006.Tiankeng:Definition and description.Speleogenesis&Evolution of Karst Aquifers,4:8.