龙脑香热带雨林附生苔藓沿宿主垂直梯度的微生境偏好及其指示作用
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摘要
该研究首次借助林冠塔吊调查了西双版纳国家级自然保护区龙脑香热带雨林样地内69棵树13个垂直高度上的附生苔藓植物,结果表明:目标样树上共记录到隶属于25科60属的90种附生苔藓,其中细鳞苔科物种数最多,占比达25.6%。13个垂直高度上共划分出三种生态类型:喜阳苔藓(散生巨树上>45 m的区域),喜阴苔藓(乔木树干上<15 m的区域),广布苔藓(广泛分布于宿主各个垂直高度上,生态位宽),并筛选出对微生境有特殊偏好的17种苔藓指示种(IndVal≥0.7,P<0.05)。随宿主垂直高度的升高,扇型和交织型的苔藓占比降低,悬垂型苔藓占比先升高后降低,细平铺型和粗平铺型的苔藓占比升高。大气湿度、水汽压、胸径以及树皮粗糙度对附生苔藓生活型的分布偏好具有显著影响。总之,沿宿主垂直高度上的附生苔藓对微环境变化在生活型和形态结构上有着不同的响应方式,而同一种生态型的苔藓群落有相似的适应机制。因此,在森林林冠生境变化的监测和管理中,对微生境具有明显偏好的附生苔藓物种可作为有效的指示材料。
Epiphytic bryophytes play significant roles in maintaining water and nutrient cycles in the forest ecosystem. However,diversity and distribution patterns of epiphytic bryophytes in forest canopies are understudied because of the limitations of canopy investigation techniques. Epiphytic bryophytes and their life-forms were investigated from 69 trees in thirteen vertical segments using a canopy crane in a tropical dipterocarp rain forest in Xishuangbanna National Nature Reserve. Canonical correlation analysis( CCA) and the species indicator method were applied to analyze the microhabitat preferences of the epiphytic bryophytes and to find indicators along the vertical gradient of the hosts. The main results were as follows: In the study area 90 species,belonging to 60 genera and 25 families,were recorded. Lejeuneaceae was the most species-rich family,representing 25.6% of all bryophyte species recorded. Three special ecological types were extracted from the thirteen vertical segments,including sun-loving bryophytes( >45 m on tree trunks in the forest canopies of emergent trees),shade-loving bryophytes( <15 m on tree trunks) and generalist bryophytes( usually distributed in various host heights with a broad niche width). Seventeen bryophyte species( Ind Val≥0.7,P<0.05) were selected as indicators for specific vertical segments. With the increase in the height sampled,the proportion of species with fan and weft life-forms decreased,the proportion of pendant life-forms increased first and then decreased,while the proportion of smooth mat and rough mat life-forms increased. The relative humidity,vapor pressure,diameter at breast height( DBH) and bark roughness had significant influences on the distribution preferences of epiphyte life forms. In conclusion,epiphytic bryophytes show different life-form compositions and morphological structures in response to changes in the micro-environment along the vertical gradient of hosts,while bryophytes assembled in the same micro-habitat usually have similar adaptation mechanisms. The epiphytic species with obvious preferences for specific microhabitats can be used as indicators during the monitoring and management of habitat changes in the forest canopy.
Epiphytic bryophytes play significant roles in maintaining water and nutrient cycles in the forest ecosystem. However,diversity and distribution patterns of epiphytic bryophytes in forest canopies are understudied because of the limitations of canopy investigation techniques. Epiphytic bryophytes and their life-forms were investigated from 69 trees in thirteen vertical segments using a canopy crane in a tropical dipterocarp rain forest in Xishuangbanna National Nature Reserve. Canonical correlation analysis( CCA) and the species indicator method were applied to analyze the microhabitat preferences of the epiphytic bryophytes and to find indicators along the vertical gradient of the hosts. The main results were as follows: In the study area 90 species,belonging to 60 genera and 25 families,were recorded. Lejeuneaceae was the most species-rich family,representing 25.6% of all bryophyte species recorded. Three special ecological types were extracted from the thirteen vertical segments,including sun-loving bryophytes( >45 m on tree trunks in the forest canopies of emergent trees),shade-loving bryophytes( <15 m on tree trunks) and generalist bryophytes( usually distributed in various host heights with a broad niche width). Seventeen bryophyte species( Ind Val≥0.7,P<0.05) were selected as indicators for specific vertical segments. With the increase in the height sampled,the proportion of species with fan and weft life-forms decreased,the proportion of pendant life-forms increased first and then decreased,while the proportion of smooth mat and rough mat life-forms increased. The relative humidity,vapor pressure,diameter at breast height( DBH) and bark roughness had significant influences on the distribution preferences of epiphyte life forms. In conclusion,epiphytic bryophytes show different life-form compositions and morphological structures in response to changes in the micro-environment along the vertical gradient of hosts,while bryophytes assembled in the same micro-habitat usually have similar adaptation mechanisms. The epiphytic species with obvious preferences for specific microhabitats can be used as indicators during the monitoring and management of habitat changes in the forest canopy.
引文
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CAO M,ZOU X,WARREN M,et al.,2006.Tropical forests of Xishuangbanna,China[J].Biotropica,38(3):306-309.
CHAZDON RL,FIELD N,1984.Light environment of tropical forest[M]//MEDINA E,MOONEY HA,VZQUEZ-YNES C.Physiological Ecology of Plants of the Wet Tropics.The Hague:Junk Publishers.
CORNELISSEN JHC,STREEGE HT,1989.Distribution and ecology of epiphytic bryophytes and lichens in dry evergreen forest of Guyana[J].J Trop Ecol,5(2):131-150.
DISLICH R,MANTOVANI W,2016.Vascular epiphyte assemblages in a brazilian atlantic forest fragment:Investigating the effect of host tree features[J].Plant Ecol,217(1):1-12.
DUFRNE M,LEGENDRE P,1997.Species assemblages and indicator species:The need for a flexible asymmetrical approach[J].Ecol Monogr,67:345-366.
ENGEMANN KB,SANDEL BJ,ENQUIST PM,et al.,2016.Patterns and drivers of plant functional group dominance across the Western Hemisphere:A macroecological re-assessment based on a massive botanical dataset[J].Bot J Linn Soc,180:141-160.
FRAHM JP,GRADSTEIN SR,1991.An altitudinal zonation of tropical rain forests using bryophytes[J].J Biogeogr,18(6):669-678.
FREY W,GOSSOW R,KRSCHNER H,1990.Verteilungsmuster von Lebensformen,wasserleitenden und wasserspeichernden Strukturen in epiphytischen Moosgesellschaften am Mt.Kinabalu(Nord-Borneo)[J].Nova Hedwigia,51:87-119.
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GLIME JM,2013.Adaptive Strategies:Growth and Life Forms.Chapt.4-5[M]//GLIME JM.Bryophyte Ecology.Volume 1.Ebook sponsored by Michigan Technological University and the International Association of Bryologists.http://digitalcommons.mtu.edu/bryophyte-ecology1/
GONZLEZ-MANCEBO JM,ROMAGUERA F,LOSADA-LIMAA,et al.,2004.Epiphytic bryophytes growing on Laurus azorica,(Seub.)Franco in three laurel forest areas in Tenerife(Canary Islands)[J].Acta Oecol,25(3):159-167.
GRADSTEIN SR,CHURCHILL SP,SALAZAR AN,2001.Guide to the bryophytes of tropical America[J].Febs Lett,26(1-2):20-24.
GRADSTEIN SR,PCS T,1989.Bryophytes[M]//LIETHH,WERGER MJA.Tropical Rain Forest Ecosystems.Amsterdam,the Netherlands:Elsevier Science Publishers:311-325.
HOLZ I,GRADSTEIN SR,2005.Cryptogamic epiphytes in primary and recovering upper montane oak forests of Costa Rica:Species richness,community composition and ecology[J].J Plant Ecol,178(1):89-109.
HOLZ I,GRADSTEIN SR,HEINRICHS J,et al.,2002.Bryophyte diversity,microhabitat differentiation and distribution of life forms in Costa Rican upper montane Quercus forest[J].Bryologist,105:334-348.
JOHANSSON D,1974.Ecology of vascular epiphytes in West African rain forest[J].Acta Pharm Suec,59:1-136.
KELLY DL,O'DONOVAN G,FEEHAN J,et al.,2004.The epiphyte communities of a montane rain forest in the andes of Venezuela:Patterns in the distribution of the flora[J].JTrop Ecol,20(6):643-666.
KRINSKY NI,1968.Chapter 5-The protective function of carotenoid pigments[M]//GIESE AC.Photophysiology:Current Topics.Elsevier Inc:123-195
KRSCHNER H,PAROLLY G,1998.Syntaxonomy of trunkepiphytic bryophyte communities of tropical rain forests.A first pantropical approach[J].Phytocoenologia,28:357-425.
KRSCHNER H,1990.Die epiphytischen Moosgesellschaften am Mt.Kinabalu(Nord-Borneo,Sabah,Malaysia)[J].Nova Hedwigia,51:1-75.
LI Q,2011.Indicator value(IndVal)method and its application[J].Chin Bull Entomol,48(2):457-462.[李巧,2011.指示值方法及其在昆虫中的应用[J].应用昆虫学报,48(2):457-462.]
LOWMAN MD,SCHOWALTER TD,2012.Plant science in forest canopies-The first 30 years of advances and challenges(1980-2010)[J].New Phytol,194:12-27.
MA KP,2015.Biodiversity monitoring in China:From CForBio to Sino BON[J].Biodivers Sci,23(1):1-2.[马克平,2015.中国生物多样性监测网络建设:从CForBio到Sino BON[J].生物多样性,23(1):1-2.]
MA KP,2015.Biodiversity monitoring relies on the integration of human observation and automatic collection of data with advanced equipment and facilities[J].Biodivers Sci,24(11):1201-1202.[马克平,2016.生物多样性监测依赖于地面人工观测与先进技术手段的有机结合[J].生物多样性,24(11):1201-1202.]
MGDEFRAU K,1982.Life-forms of bryophytes[M]//SMITH AJE.Bryophyte Ecology[M].Chapman and Hall,London:45-58.
MALE TD,ROBERTS GE,2005.Host associations of the strangler fig Ficus watkinsiana in a subtropical Queensland rain forest[J].Austral Ecol,30:229-236.
MITCHELL AW,SECOY K,JACKSON T,2002.The global canopy handbook:Techniques of access and study forest roof[M].Oxford,UK:Global Canopy Programme.
MITTERMEIER RA,ROBLES GP,HOFFMANN M,et al.,2004.Hotspots revisited:Earth’s biologically richest and most endangered terrestrial ecoregions[M].Mexico City:CEMEX.
NAKAMURA A,KITCHING RL,CAO M,et al.,2017.Forests and their canopies:Achievements and horizons in canopy science[J].Trend Ecol Evol,32(6):438-451.
OKSANEN J,BLANCHET FG,FRIENDLY M,et al.,2017.Vegan:Community Ecology Package.R package version 2.4-4.https://CRAN.R-project.org/package=vegan
OLIVEIRA HCD,OLIVEIRA SMD,OLIVEIRA HCD,et al.,2016.Vertical distribution of epiphytic bryophytes in Atlantic forest fragments in northeastern Brazil[J].Acta Bot Brasi:30(ahead).
RAES N,CANNON CH,HIJMANS RJ,et al.,2014.Historical distribution of sundaland's dipterocarp rainforests at quaternary glacial maxima[J].Proc Natl Acad Sci USA,111(47):16790-16795.
ROBERTS DW,2016.Labdsv:Ordination and multivariate analysis for ecology.R package version 1.8-0.https://cran.r-project.org/web/packages/labdsv
SCOTT GAM,1982.Desert bryophytes[M]//SMITH AJE.Bryophyte Ecology.London:Chapman and Hall:105-122.
SHI XM,SONG L,LIU WY,et al.,2017.Epiphytic bryophytes as bio-indicators of atmospheric nitrogen deposition in a subtropical montane cloud forest:Response patterns,mechanism,and critical load[J].Environ Poll,229(X):932-941.
SLACK NG,1976.Host specificity of bryophytic epiphytes in eastern North America(Geography and Ecology of Bryophytes)[J].J Hatt Bot Lab,41:107-132.
SMITH AJE,1982.Bryophytes and Epiliths in Bryophyte Ecology[M]//SMITH AJE.Bryophyte Ecology.London:Chapman and Hall:1-14.
SONG L,LIU WY,MA WZ,et al.,2011.Bole epiphytic bryophytes on Lithocarpus xylocarpus,(Kurz)Markgr.in the Ailao Mountains,SW China[J].Ecol Res,26(2):351-363.
SPORN SG,BOS MM,KESSLER M,et al.,2010.Vertical distribution of epiphytic bryophytes in an Indonesian rainforest[J].Biodivers Conserv,19(3):745-760.
SPORN SG,BOS MM,HOFFSTTTER-MNCHEBERG M,et al.,2009.Microclimate determines community composition but not richness of epiphytic understorey bryophytes of rainforest and cacao agroforests in Indonesia[J].Funct Plant Biol,36:171-179.
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