五台山林线附近优势草本植物的分布格局、过程与尺度分析
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摘要
林线是气候变化敏感指示区,林窗和林缘是草本植物进入的重要窗口。研究林线过渡带林窗和林缘生境内草本物种的格局有利于揭示这两种生境内重要的生态过程。本文选取五台山自然保护区林线过渡带内的林窗(海拔2350 m)和林缘(海拔2300 m)为样地(5 m×5 m),每个样地设置25个1 m×1 m样方,分别记录样方内6个优势物种(光稃茅香(Hierochloe glabra)、小花草玉梅(Anemone rivularis)、平车前(Plantago depressa)、珠芽蓼(Polygonum viviparum)、三叶委陵菜(Potentilla freyniana)、蒲公英(Taraxacum mongolicum))位置信息。运用完全空间随机过程(CSR)、异质性泊松过程(HP)、泊松聚块过程(PC)零模型在不同尺度下进行格局模拟分析。结果如下:(1)林窗内6个物种的聚集分布的尺度范围都分别大于林缘,在两个生境内珠芽蓼和小花草玉梅聚集尺度较宽,蒲公英和平车前聚集尺度较窄,最常见的聚集尺度在0～40、100～140和200～230 cm。(2)异质性泊松过程模拟及检验结果表明:在林窗和林缘,微生境异质性对具有广适性的5个物种的格局影响不显著,而无性繁殖较发达的光稃茅香的格局在林窗和林缘生境内受微生境影响显著。(3)泊松聚块过程模拟及检验结果表明:林缘的物种格局显著受到种子扩散过程的影响。在0～50 cm的尺度范围内种子扩散过程受重力影响分布在母体周围。在大于100 cm尺度上,种子特殊的传播方式或特殊种子附属结构影响种子扩散的距离。此外,在风力帮助下,特殊结构的物种分布有可能达到250 cm。
Treeline is sensitive to climate change. Forest gap and edge are important windows for the invasion of herbaceous species. Research on distribution patterns of herbaceous species in forest gaps and forest edges of treeline ecotones could help explain important ecological processes.We selected a forest gap( elevation: 2350 m) and a forest edge( elevation: 2300 m) as plots( 5 m×5 m) in a treeline ecotone of Wutai Natural Reserve. Twenty-five quadrats( 1 m×1 m)were set up in each of both plots,within which the locations of six dominant species( Hierochloe glabra,Anemone rivularis,Plantago depressa,Polygonum viviparum,Potentilla freyniana,and Taraxacum mongolicum) were recorded. Complete spatial random process( CSR),heterogeneous Poisson process( HP),and Poisson cluster( PC) were used to simulate the spatial patterns of the six species at different scales. The results showed that:( 1) Aggregated sizes of the six species in forest gaps were broader than those in the forest edge,with those of Polygonum viviparum and Anemone rivularis were broader than other species in both plots. The aggregated sizes of Taraxacum mongolicum and Plantago depressa were narrower than other species in both plots. The most common aggregation occurred on 0-40,100-140 and 200-230 cm.( 2) For both forest gap and edge,HP simulation and significance tests showed that micro-habitat differences between plots had no significant effect on species distribution with broad ecological amplitude. However,the distribution of Hierochloe glabra was significantly affected by micro-habitat differences in forest gap and edge due to ramet growth.( 3) Results from PC simulations and significance tests showed that seed dispersal significantly affected species distribution in the forest edge. Seed dispersal around maternal individuals often occurred at 0-50 cm scales as affected by gravity. At the scale of >100 cm,species with special dispersal modes or structures affected their dispersal distance. Additionally,species with special structures could reach 250 cm with the help of strong winds.
Treeline is sensitive to climate change. Forest gap and edge are important windows for the invasion of herbaceous species. Research on distribution patterns of herbaceous species in forest gaps and forest edges of treeline ecotones could help explain important ecological processes.We selected a forest gap( elevation: 2350 m) and a forest edge( elevation: 2300 m) as plots( 5 m×5 m) in a treeline ecotone of Wutai Natural Reserve. Twenty-five quadrats( 1 m×1 m)were set up in each of both plots,within which the locations of six dominant species( Hierochloe glabra,Anemone rivularis,Plantago depressa,Polygonum viviparum,Potentilla freyniana,and Taraxacum mongolicum) were recorded. Complete spatial random process( CSR),heterogeneous Poisson process( HP),and Poisson cluster( PC) were used to simulate the spatial patterns of the six species at different scales. The results showed that:( 1) Aggregated sizes of the six species in forest gaps were broader than those in the forest edge,with those of Polygonum viviparum and Anemone rivularis were broader than other species in both plots. The aggregated sizes of Taraxacum mongolicum and Plantago depressa were narrower than other species in both plots. The most common aggregation occurred on 0-40,100-140 and 200-230 cm.( 2) For both forest gap and edge,HP simulation and significance tests showed that micro-habitat differences between plots had no significant effect on species distribution with broad ecological amplitude. However,the distribution of Hierochloe glabra was significantly affected by micro-habitat differences in forest gap and edge due to ramet growth.( 3) Results from PC simulations and significance tests showed that seed dispersal significantly affected species distribution in the forest edge. Seed dispersal around maternal individuals often occurred at 0-50 cm scales as affected by gravity. At the scale of >100 cm,species with special dispersal modes or structures affected their dispersal distance. Additionally,species with special structures could reach 250 cm with the help of strong winds.
引文
曹燕丽,崔海亭,刘鸿雁,等.2001.五台山高山带景观的遥感分析.地理学报,56(3):297-306.
常学礼,吕世海,冯朝阳,等.2015.地形对草甸草原植被生产力分布格局的影响.生态学报,35(10):3339-3348.
崔宁洁,刘洋,张健,等.2014.林窗对马尾松人工林植物多样性的影响.应用与环境生物学报,20(1):8-14.
崔本义,朱世忠,曳红玉,等.2011.五台山植物资源.太原:山西科学技术出版社:3-10.
陈文年,吴宁,罗鹏,等.2003.岷江上游林缘祁连山圆柏群落的物种多样性及乔木种群的分布格局.应用与环境生物学报,9(3):221-225.
段文标,王丽霞,陈立新,等.2013.红松阔叶混交林林隙大小及光照对草本植物的影响.应用生态学报,24(3):614-620.
范邓妹,杨永平.2009.不同生境下珠芽蓼(蓼科)的繁殖策略比较.云南植物研究,31(2):153-157.
葛晓改,周本智,王刚,等.2014.雪灾干扰下林窗对木荷幼苗更新的影响.林业科学研究,27(4):529-535.
何玉惠,赵哈林,刘新平,等.2008.不同类型沙地狗尾草的生长特征及生物量分配,生态学杂志,27(4):504-508.
贾美玉,李雪华,吴忠铉,等.2016.科尔沁沙地丘间低地不同生活型物种格局与共存机制.生态学杂志,35(1):111-117.
贾美玉,李雪华,吴忠铉,等.2015.科尔沁沙地流动沙丘3种常见植物的空间分布格局与异速生长.应用生态学报,26(10):2953-2960.
路远,龙瑞军,林丽.2007.高寒珠芽蓼草甸植被生物量的季节动态特征.草原与草坪,(1):1-4.
刘辉,宋会兴,杨万勤,等.2015.油樟幼苗对马尾松林窗面积的光合响应特征.生态学报,35(12):4089-4096.
刘秀丽,张勃,任媛,等.2015.五台山地区草地生态系统服务价值估算.干旱区资源与环境,29(5):24-29.
刘鸿雁,曹艳丽,田军,等.2003.山西五台山高山林线的植被景观.植物生态学报,27(2):263-269.
缪宁,刘世荣,史作民,等.2011.青藏高原东缘林线杜鹃-岷江冷杉原始林的空间格局.生态学报,31(1):1-9.
唐振兴,何志斌,刘鹄.2012.祁连山中段林缘土壤水热特征及其对气象要素的响应.生态学报,32(4):52-61.
谭芮,李伟,杨宇明,等.2013.不同干扰条件下纳帕海湿地植物群落优势种群的生态位.湖泊科学,25(5):681-687.
田丽,周新宇,孙航,等.2009.7种委陵菜属植物叶表皮微形态的研究.草业科学,26(2):53-56.
武玉珍,张峰.2006.山西桑干河流域湿地植被优势种群分布格局研究.植物研究,26(6):735-741.
王晓雨,于大炮,周莉,等.2015.长白山北坡林线岳桦种群空间分布格局.生态学报,35(1):116-124.
向春玲,张金屯.2013.东灵山亚高山草甸优势种的点格局分析.草业科学,30(3):317-321.
杨开军,熊莉,杨万勤,等.2016.林窗对川西亚高山云杉人工林土壤呼吸的影响.应用与环境生物学报,22(3):505-509.
张金屯.1998.植物种群空间分布的点格局分析.植物生态学报,22(4):344-349.
张金屯,孟东平.2004.芦芽山华北落叶松林不同龄级立木的点格局分析.生态学报,24(1):35-40.
张金屯.2011.数量生态学(第二版).北京:科学出版社:277-280.
张大勇.2000.理论生态学研究.北京:高等教育出版社:170-172.
张晶然.2016.施肥对青藏高原高寒草甸植物叶片氮磷重吸收的影响(硕士学位论文).兰州:兰州大学.
周桂玲.2013.稃的有无对大赖草种子扩散与萌发特性的影响.植物资源与环境学报,22(4):50-54.
周晓岭.2012.异质生境下克隆植物羊草和光稃茅香的空间拓展策略(硕士学位论文).长春:东北师范大学.
Bullock JM,Moy IL,Coulson SJ,et al.2003.Habitat-specific dispersal:Environmental effects on the mechanisms and patterns of seed movement in a grassland herb Rhinanthus minor.Ecography,26:692-704.
Browning DM,Franklin J,Archer SR,et al.2014.Spatial patterns of grassland-shrubland state transitions:A 74-year record on grazed and protected areas.Ecological Applications,24:1421-1433.
Bullock JM,Kenward RE,Hails RS.2002.Plant dispersal and colonization processes at local and landscape scales//Bullock JM,Moy IL,Pywell RF,eds.Dispersal Ecology.Oxford:Blackwell Publishing Company:279-302.
Cain ML,Damman H,Muir A.1998.Seed dispersal and the Holocene migration of woodland herbs.Ecological Monographs,68:325-347.
Carvalhais N,Forkel M,Khomik M,et al.2014.Global covariation of carbon turnover times with climate in terrestrial ecosystems.Nature,514:213-217.
Dullinger S,Dirnb9ck T,Grabherr G.2004.Modelling climate change-driven treeline shifts:Relative effects of temperature increase,dispersal and invasibility.Journal of Ecology,92:241-252.
Gilliam F.2007.The ecological significance of the herbaceous layer in temperate forest ecosystems.Bio Science,57:845-858.
Hattenschwiler S,Smith WK.1999.Seedling occurrence in alpine treeline conifers:A case study from the central Rocky Mountains,USA.Acta Oecologica,20:219-224.
Hughes NM,Johnson DM,Akhalkatsi M,et al.2009.Characterizing Betula litwinowii seedling microsites at the alpinetreeline ecotone,Central Greater Caucasus Mountains,Georgia.Arctic,Antarctic,and Alpine Research,41:112-118.
Janzen DH.1984.Dispersal of small seeds by big herbivores:Foliage is the fruit.The American Naturalist,123:338-353.
Kullman L.2014.Treeline(Pinus sylvestris)landscape evolution in the Swedish Scandes:A 40-year demographic effort viewed in a broader temporal context.Norsk Geografisk Tidsskrift-Norwegian Journal of Geography,68:155-167.
Liu HY,Tang ZY,Dai JH,et al.2002.Larch timberline and its development in North China.Mountain Research and Development,22:359-367.
Loydi A,Lohse K,Otte A,et al.2013.Distribution and effects of tree leaf litter on vegetation composition and biomass in a forest-grassland ecotone.Journal of Plant Ecology,7:264-275.
Nilsson MC,Wardle DA.2005.Understory vegetation as a forest ecosystem driver:Evidence from the northern Swedish boreal forest.Frontiers in Ecology and the Environment,3:421-428.
Sherman RE,Fahey TJ,Battles JJ.2000.Small-scale disturbance and regeneration dynamics in a neotropical mangrove forest.Journal of Ecology,88:165-178.
Shen GC,Yu MJ,Hu XS,et al.2009.Species-area relationships explained by the joint effects of dispersal limitation and habitat heterogeneity.Ecology,90:3033-3041.
Wang XG,Wiegand T,Hao ZQ,et al.2010.Species associations in an old-growth temperate forest in north-eastern China.Journal of Ecology,98:674-686.
Wang XG,Wiegand T,Wolf A,et al.2011.Spatial patterns of tree species richness in two temperate forests.Journal of Ecology,99:1382-1393.
Wiegand T,Moloney KA.2004.Rings,circles,and null-models for point pattern analysis in ecology.Oikos,104:209-229.
Wittkuhn RS.2010.Wind-aided seed dispersal of perennial Veld Grass(Ehrharta calycina):Implications for restoration in weedy urban bushland remnants.Ecological Management&Restoration,11:148-150.
常学礼,吕世海,冯朝阳,等.2015.地形对草甸草原植被生产力分布格局的影响.生态学报,35(10):3339-3348.
崔宁洁,刘洋,张健,等.2014.林窗对马尾松人工林植物多样性的影响.应用与环境生物学报,20(1):8-14.
崔本义,朱世忠,曳红玉,等.2011.五台山植物资源.太原:山西科学技术出版社:3-10.
陈文年,吴宁,罗鹏,等.2003.岷江上游林缘祁连山圆柏群落的物种多样性及乔木种群的分布格局.应用与环境生物学报,9(3):221-225.
段文标,王丽霞,陈立新,等.2013.红松阔叶混交林林隙大小及光照对草本植物的影响.应用生态学报,24(3):614-620.
范邓妹,杨永平.2009.不同生境下珠芽蓼(蓼科)的繁殖策略比较.云南植物研究,31(2):153-157.
葛晓改,周本智,王刚,等.2014.雪灾干扰下林窗对木荷幼苗更新的影响.林业科学研究,27(4):529-535.
何玉惠,赵哈林,刘新平,等.2008.不同类型沙地狗尾草的生长特征及生物量分配,生态学杂志,27(4):504-508.
贾美玉,李雪华,吴忠铉,等.2016.科尔沁沙地丘间低地不同生活型物种格局与共存机制.生态学杂志,35(1):111-117.
贾美玉,李雪华,吴忠铉,等.2015.科尔沁沙地流动沙丘3种常见植物的空间分布格局与异速生长.应用生态学报,26(10):2953-2960.
路远,龙瑞军,林丽.2007.高寒珠芽蓼草甸植被生物量的季节动态特征.草原与草坪,(1):1-4.
刘辉,宋会兴,杨万勤,等.2015.油樟幼苗对马尾松林窗面积的光合响应特征.生态学报,35(12):4089-4096.
刘秀丽,张勃,任媛,等.2015.五台山地区草地生态系统服务价值估算.干旱区资源与环境,29(5):24-29.
刘鸿雁,曹艳丽,田军,等.2003.山西五台山高山林线的植被景观.植物生态学报,27(2):263-269.
缪宁,刘世荣,史作民,等.2011.青藏高原东缘林线杜鹃-岷江冷杉原始林的空间格局.生态学报,31(1):1-9.
唐振兴,何志斌,刘鹄.2012.祁连山中段林缘土壤水热特征及其对气象要素的响应.生态学报,32(4):52-61.
谭芮,李伟,杨宇明,等.2013.不同干扰条件下纳帕海湿地植物群落优势种群的生态位.湖泊科学,25(5):681-687.
田丽,周新宇,孙航,等.2009.7种委陵菜属植物叶表皮微形态的研究.草业科学,26(2):53-56.
武玉珍,张峰.2006.山西桑干河流域湿地植被优势种群分布格局研究.植物研究,26(6):735-741.
王晓雨,于大炮,周莉,等.2015.长白山北坡林线岳桦种群空间分布格局.生态学报,35(1):116-124.
向春玲,张金屯.2013.东灵山亚高山草甸优势种的点格局分析.草业科学,30(3):317-321.
杨开军,熊莉,杨万勤,等.2016.林窗对川西亚高山云杉人工林土壤呼吸的影响.应用与环境生物学报,22(3):505-509.
张金屯.1998.植物种群空间分布的点格局分析.植物生态学报,22(4):344-349.
张金屯,孟东平.2004.芦芽山华北落叶松林不同龄级立木的点格局分析.生态学报,24(1):35-40.
张金屯.2011.数量生态学(第二版).北京:科学出版社:277-280.
张大勇.2000.理论生态学研究.北京:高等教育出版社:170-172.
张晶然.2016.施肥对青藏高原高寒草甸植物叶片氮磷重吸收的影响(硕士学位论文).兰州:兰州大学.
周桂玲.2013.稃的有无对大赖草种子扩散与萌发特性的影响.植物资源与环境学报,22(4):50-54.
周晓岭.2012.异质生境下克隆植物羊草和光稃茅香的空间拓展策略(硕士学位论文).长春:东北师范大学.
Bullock JM,Moy IL,Coulson SJ,et al.2003.Habitat-specific dispersal:Environmental effects on the mechanisms and patterns of seed movement in a grassland herb Rhinanthus minor.Ecography,26:692-704.
Browning DM,Franklin J,Archer SR,et al.2014.Spatial patterns of grassland-shrubland state transitions:A 74-year record on grazed and protected areas.Ecological Applications,24:1421-1433.
Bullock JM,Kenward RE,Hails RS.2002.Plant dispersal and colonization processes at local and landscape scales//Bullock JM,Moy IL,Pywell RF,eds.Dispersal Ecology.Oxford:Blackwell Publishing Company:279-302.
Cain ML,Damman H,Muir A.1998.Seed dispersal and the Holocene migration of woodland herbs.Ecological Monographs,68:325-347.
Carvalhais N,Forkel M,Khomik M,et al.2014.Global covariation of carbon turnover times with climate in terrestrial ecosystems.Nature,514:213-217.
Dullinger S,Dirnb9ck T,Grabherr G.2004.Modelling climate change-driven treeline shifts:Relative effects of temperature increase,dispersal and invasibility.Journal of Ecology,92:241-252.
Gilliam F.2007.The ecological significance of the herbaceous layer in temperate forest ecosystems.Bio Science,57:845-858.
Hattenschwiler S,Smith WK.1999.Seedling occurrence in alpine treeline conifers:A case study from the central Rocky Mountains,USA.Acta Oecologica,20:219-224.
Hughes NM,Johnson DM,Akhalkatsi M,et al.2009.Characterizing Betula litwinowii seedling microsites at the alpinetreeline ecotone,Central Greater Caucasus Mountains,Georgia.Arctic,Antarctic,and Alpine Research,41:112-118.
Janzen DH.1984.Dispersal of small seeds by big herbivores:Foliage is the fruit.The American Naturalist,123:338-353.
Kullman L.2014.Treeline(Pinus sylvestris)landscape evolution in the Swedish Scandes:A 40-year demographic effort viewed in a broader temporal context.Norsk Geografisk Tidsskrift-Norwegian Journal of Geography,68:155-167.
Liu HY,Tang ZY,Dai JH,et al.2002.Larch timberline and its development in North China.Mountain Research and Development,22:359-367.
Loydi A,Lohse K,Otte A,et al.2013.Distribution and effects of tree leaf litter on vegetation composition and biomass in a forest-grassland ecotone.Journal of Plant Ecology,7:264-275.
Nilsson MC,Wardle DA.2005.Understory vegetation as a forest ecosystem driver:Evidence from the northern Swedish boreal forest.Frontiers in Ecology and the Environment,3:421-428.
Sherman RE,Fahey TJ,Battles JJ.2000.Small-scale disturbance and regeneration dynamics in a neotropical mangrove forest.Journal of Ecology,88:165-178.
Shen GC,Yu MJ,Hu XS,et al.2009.Species-area relationships explained by the joint effects of dispersal limitation and habitat heterogeneity.Ecology,90:3033-3041.
Wang XG,Wiegand T,Hao ZQ,et al.2010.Species associations in an old-growth temperate forest in north-eastern China.Journal of Ecology,98:674-686.
Wang XG,Wiegand T,Wolf A,et al.2011.Spatial patterns of tree species richness in two temperate forests.Journal of Ecology,99:1382-1393.
Wiegand T,Moloney KA.2004.Rings,circles,and null-models for point pattern analysis in ecology.Oikos,104:209-229.
Wittkuhn RS.2010.Wind-aided seed dispersal of perennial Veld Grass(Ehrharta calycina):Implications for restoration in weedy urban bushland remnants.Ecological Management&Restoration,11:148-150.
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