海南尖峰岭热带山地雨林60 ha大样地木本植物性别系统数量特征
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摘要
热带山地雨林中植物不同性别系统的数量和空间分布特征如何?是否受自然环境条件影响?这些问题的回答有助于更深入理解群落物种多样性是如何形成的。本文以海南尖峰岭热带山地雨林60 ha大样地中胸径大于1 cm的木本植物为研究对象,描述了样地内木本植物性别系统的数量特征,分析了雌雄异株植物空间分布与地形因子的相关关系。结果表明:大样地内289种木本种子植物中有两性花植物176种,单性花植物113种;在单性花植物中,51种为雌雄同株, 62种为雌雄异株。单性花植物植株数占所有植株总数的36.2%;雌雄异株植物植株数占所有植株总数的21.5%,占单性花植物植株数的59.5%。在20 m×20 m样方的空间尺度,雌雄异株植物呈现聚集分布的物种共有31种。以20 m×20 m样方为分析单元,雌雄异株植物种类的性别比例与海拔、凹凸度和坡度呈弱正相关;个体相对多度仅和海拔呈弱正相关,与坡度和凹凸度无相关;而3个性别系统多样性指数与海拔、凹凸度和坡度均无显著关联。可见雌雄异株植物在海南热带山地雨林中占据了较大的比例,但大部分种类种群较小,其数量分布特征与地形因素紧密相关,雌雄异株植物的存在对群落物种多样性的形成有较大贡献。
How plants with different sexual systems grow and distribute in the community? Are they influenced by environmental factors? The answer to these questions is helpful to understand how species diversity is shaped in the plant community. This study summarized the numerical characteristics of sexual systems and the relationship between the distribution of dioecious plants and the topographical factors. Results showed that of the 289 woody seed plants recorded in the 60 ha plot, 176 species are bisexual plants and 113 species are unisexual plants. Of the 113 unisexual species, 51 are monoecious and 62 are dioecious plants. The abundance of unisexual plants accounts for 36.2% of all individuals recorded in the whole plot. The dioecious plants account for 21.5% of all individuals recorded and 59.5% of all individuals of unisexual plants in the plot. At the 20 m × 20 m quadrat spatial scale, 31 dioecious species showed aggregated distribution. Proportion of number of dioecious species of all species showed weak and positive correlation with altitude, convex and slope, but relative abundance of dioecious plants only showed weak and positive correlation with altitude. Furthermore, three diversity indices of plant sexual system did not show significant correlation with altitude, convex and slope. In sum, dioecious plants occupy a large proportion but most species only have relatively small population in the tropical mountain rain forest in Hainan, which are closely correlated with the topographical factors by the changes of numerical characteristics. The existence of dioecious plants has important contribution to the formation of species diversity in the plant community.
How plants with different sexual systems grow and distribute in the community? Are they influenced by environmental factors? The answer to these questions is helpful to understand how species diversity is shaped in the plant community. This study summarized the numerical characteristics of sexual systems and the relationship between the distribution of dioecious plants and the topographical factors. Results showed that of the 289 woody seed plants recorded in the 60 ha plot, 176 species are bisexual plants and 113 species are unisexual plants. Of the 113 unisexual species, 51 are monoecious and 62 are dioecious plants. The abundance of unisexual plants accounts for 36.2% of all individuals recorded in the whole plot. The dioecious plants account for 21.5% of all individuals recorded and 59.5% of all individuals of unisexual plants in the plot. At the 20 m × 20 m quadrat spatial scale, 31 dioecious species showed aggregated distribution. Proportion of number of dioecious species of all species showed weak and positive correlation with altitude, convex and slope, but relative abundance of dioecious plants only showed weak and positive correlation with altitude. Furthermore, three diversity indices of plant sexual system did not show significant correlation with altitude, convex and slope. In sum, dioecious plants occupy a large proportion but most species only have relatively small population in the tropical mountain rain forest in Hainan, which are closely correlated with the topographical factors by the changes of numerical characteristics. The existence of dioecious plants has important contribution to the formation of species diversity in the plant community.
引文
Chen J,Flemming TH,Zhang L,Wang H,Liu Y(2004)Patterns of fruit traits in a tropical rainforest in Xishuangbanna,SW China.Acta Oecologica,26,157-164.
Chen J,Han QQ,Duan BL,Korpelainen H,Li CY(2017)Sex-specific competition differently regulates ecophysiological responses and phytoremediation of Populus cathayana under Pb stress.Plant and Soil,421,203-218.
Chen XS(2008)Community Level Plant Reproductive Traits of Tropical Rain Forest in Xishuangbanna,SW China.PhDdissertation,Xishuangbanna Tropical Botanical Garden,Chinese Academy of Sciences,Xishuangbanna,Mengla,Yunnan.(in Chinese with English abstract)[陈心胜(2008)西双版纳热带雨林植物群落的繁殖生物学特性研究.博士学位论文,中国科学院西双版纳热带植物园,云南勐腊.]
Chen XS,Li QJ(2008)Sexual systems and ecological correlates in an azonal tropical forests,SW China.Biotropica,40,160-167.
Darwin C(1877)The Different Forms of Flowers on Plants of the Same Species.John Murray,London.
Diggle PJ(1983)Statistical Analysis of Spatial Point Patterns.Academic Press,New York.
Du RQ(2009)Biostatistics.Higher Education Press,Beijing.(in Chinese)[杜荣骞(2009)生物统计学.高等教育出版社,北京.]
Graff P,Rositano F,Aguiar MR(2013)Changes in sex ratios of a dioecious grass with grazing intensity:The interplay between gender traits,neighbour interactions and spatial patterns.Journal of Ecology,101,1146-1157.
Heikrujam M,Sharma K,Prasad M,Agrawal V(2015)Review on different mechanisms of sex determination and sexlinked molecular markers in dioecious crops-A current update.Euphytica,201,161-194.
Henry IM,Akagi T,Tao R,Comai L(2018)One hundred ways to invent the sexes:Theoretical and observed paths to dioecy in plants.Annual Review of Plant Biology,69,553-575.
Huang SQ,Guo YH(2000)Progress in pollination biology.Chinese Science Bulletin,45,225-237.(in Chinese with English abstract)[黄双全,郭友好(2000)传粉生物学的研究进展.科学通报,45,225-237.]
Hultine KR,Grady KC,Wood TE,Shuster SM,Stella JC,Whitham TG(2016)Climate change perils for dioecious plant species.Nature Plants,2,16109.
James DM(1991)Botany:An Introduction to Plant Biology.Saunders College Publishing,Austin.
Jiang YX,Lu JP(1991)Ecosystem of Tropical Forest of Jianfengling Mountain,Hainan Island,China.Science Press,Beijing.(in Chinese)[蒋有绪,卢俊培(1991)中国海南岛尖峰岭热带林生态系统.科学出版社,北京.]
Jing SW,Coley PD(1990)Dioecy and herbivory:The effect of growth rate on plant defense in Acer negundo.Oikos,58,369-377.
Juvany M,Munné-Bosch S(2015)Sex-related differences in stress tolerance in dioecious plants:A critical appraisal in a physiological context.Journal of Experimental Botany,66,6083-6092.
Li TH,Jiang J,Chen JM,Fan SB(2004)The sexual polymorphism of seed plant.Journal of Northeast Forestry University,32,48-52.(in Chinese with English abstract)[李同华,姜静,陈建名,范士波(2004)种子植物性别的多态性.东北林业大学学报,32,48-52.]
Li YD,Chen BF,Zhou GY,Wu ZM,Zeng QB,Luo TS,Huang SN,Xie MD,Huang Q(2002)Research and Conservation of Tropical Forest and the Biodiversity:A Special Reference to Hainan Island,China.China Forestry Publishing House,Beijing.(in Chinese)[李意德,陈步峰,周光益,吴仲民,曾庆波,骆土寿,黄世能,谢明东,黄全(2002)中国海南岛热带森林及其生物多样性保护研究.中国林业出版社,北京.]
Li YD,Xu H,Luo TS,Chen DX,Lin MX(2012)Bio-Checklist of Jianfengling Station.China Agricultural Press,Beijing.(in Chinese)[李意德,许涵,骆土寿,陈德祥,林明献(2012)尖峰岭生态站生物物种数据集.中国农业出版社,北京.]
Long R(2011)Study on Community Structure of Deciduous Broad-Leaf Forests and Their Plant Sexual System Diversity in Dalaoling,Hubei Province.PhD Dissertation,Beijing Forestry University,Beijing.(in Chinese with English abstract)[龙茹(2011)湖北大老岭落叶阔叶林群落结构和植物性系统多样性研究.博士学位论文,北京林业大学,北京.]
Long R,Shang C,Qu S,Zhang ZX(2011)Distribution pattern of plant sexual system diversity in Populus lasiocarpa Oliv.community.Journal of Beijing Forestry University,33,34-41.(in Chinese with English abstract)[龙茹,尚策,曲上,张志翔(2011)大叶杨群落植物性系统多样性的分布格局.北京林业大学学报,33,34-41.]
Machado LC,Vlopes A,Sazima M(2006)Plant sexual systems and review of the breeding system studies in the Caatinga,a Brazilian tropical dry forest.Annals of Botany,97,277-287.
Maxime RM,Pierre-Olivier C(2015)High incidence of dioecy in young successional tropical forests.Journal of Ecology,103,725-732.
McIntosh RP(1967)The continuum concept of vegetation.The Botanical Review,33,130-187.
Pannell JR(2017)Plant sex determination.Current Biology,27,191-197.
Pielou EC(1975)Ecological Diversity.John Wiley&Sons,New York.
Ramirez N(2005)Plant sexual systems,dichogamy,and herkogamy in the Venezuelan Central Plain.Flora Jena,200,30-48.
Ramirez N,Brito Y(1990)Reproductive biology of a tropical palm swamp community in the Venezuelan LIanos.American Journal of Botany,77,1260-1271.
Renner(2014)The relative and absolute frequencies of angiosperm sexual systems:Dioecy,monoecy,gynodioecy,and an updated online database.American Journal of Botany,101,1588-1596.
Robinson J(2008)The Evolution of Flower Size and Flowering Behaviour in Plants:The Role of Pollination and Pre-dispersal Seed Predation.Master dissertation,University of Southampton,UK.
Shannon CE(1949)Communication theory of secrecy systems.The Bell System Technical Journal,28,656-715.
Simpson EH(1949)Measurement of diversity.Nature,163,688.
Tao J,Zang RG,Li YD,Mao PL,Lin MX(2011)Characteristic of plant functional groups based on ecophysiological traits in a tropical montane rain forest of Hainan Island,South China.Scientia Silvae Sinicae,47(8),14-18.(in Chinese with English abstract)[陶晶,臧润国,李意德,毛培利,林明献(2011)海南尖峰岭热带山地雨林生理生态功能群特征.林业科学,47(8),14-18.]
Xu H,Li YD,Lin MX,Wu JH,Luo TS,Zhou Z,Chen DX,Yang H,Li GJ,Liu SR(2015a)Community characteristics of a 60 ha dynamics plot in the tropical montane rain forest in Jianfengling,Hainan Island.Biodiversity Science,23,192-201.(in Chinese with English abstract)[许涵,李意德,林明献,吴建辉,骆土寿,周璋,陈德祥,杨怀,李广建,刘世荣(2015a)海南尖峰岭热带山地雨林60 ha动态监测样地群落结构特征.生物多样性,23,192-201.]
Xu H,Li YD,Luo TS,Chen DX,Lin MX,Wu JH,Li YP,Yang H,Zhou Z(2015b)Jianfengling Tropical Mountain Rain Forest Dynamic Plot:Community Characteristics,Tree Species and Their Distribution Patterns.China Forestry Publishing House,Beijing.(in Chinese)[许涵,李意德,骆土寿,陈德祥,林明献,吴建辉,李艳朋,杨怀,周璋(2015b)海南尖峰岭热带山地雨林--群落特征、树种及其分布格局.中国林业出版社,北京.]
Xu H,Li YD,Luo TS,Lin MX,Chen DX,Mo JH,Luo W,Huang H(2008)Influence of typhoon Damrey on the tropical montane rain forest community in Jianfengling,Hainan Island,China.Chinese Journal of Plant Ecology,32,1323-1334.(in Chinese with English abstract)[许涵,李意德,骆土寿,林明献,陈德祥,莫锦华,罗文,黄豪(2008)达维台风对海南尖峰岭热带山地雨林群落的影响.植物生态学报,32,1323-1334.]
Xu X,Yang F,Yin CY,Li CY(2007)Research advances in sex-specific responses of dioecious plants to environmental stresses.Chinese Journal of Applied Ecology,18,2626-2631.(in Chinese with English abstract)[胥晓,杨帆,尹春英,李春阳(2007)雌雄异株植物对环境胁迫响应的性别差异研究进展.应用生态学报,18,2626-2631.]
Yamakura T,Kanzaki M,Itoh A,Ohkubo T,Ogino K,Chai EOK,Lee HS,Ashton PS(1995)Topography of a large-scale research plot established within a tropical rain forest at Lambir,Sarawak.Tropics,5,41-56.
Yan LH(2007)Sexual system and environmental adaptability of vines in Hupingshan Mountain,Hunan Province.Journal of Northeast Forestry University,35,35-36,39.(in Chinese with English abstract)[颜立红(2007)湖南壶瓶山藤本植物的有性系统及其环境适应性.东北林业大学学报,35,35-36,39.]
Yin CY,Li CY(2007)Gender differences of dioecious plants related sex ratio:Recent advances and future prospects.Chinese Journal of Applied Environmental Biology,13,419-425.(in Chinese with English abstract)[尹春英,李春阳(2007)雌雄异株植物与性别比例有关的性别差异研究现状与展望.应用与环境生物学报,13,419-425.]
Zeng QB,Li YD,Chen BF,Wu ZM,Zhou GY(1997)Research and Management of Tropical Forest Ecosystem.China Forestry Publishing House,Beijing.(in Chinese)[曾庆波,李意德,陈步峰,吴仲民,周光益(1997)热带森林生态系统研究与管理.中国林业出版社,北京.]
Zhang DY(2004)Plant Life-History Evolution and Reproductive Ecology.Science Press,Beijing.(in Chinese)[张大勇(2004)植物生活史进化与繁殖生态学.科学出版社,北京.]
Chen J,Han QQ,Duan BL,Korpelainen H,Li CY(2017)Sex-specific competition differently regulates ecophysiological responses and phytoremediation of Populus cathayana under Pb stress.Plant and Soil,421,203-218.
Chen XS(2008)Community Level Plant Reproductive Traits of Tropical Rain Forest in Xishuangbanna,SW China.PhDdissertation,Xishuangbanna Tropical Botanical Garden,Chinese Academy of Sciences,Xishuangbanna,Mengla,Yunnan.(in Chinese with English abstract)[陈心胜(2008)西双版纳热带雨林植物群落的繁殖生物学特性研究.博士学位论文,中国科学院西双版纳热带植物园,云南勐腊.]
Chen XS,Li QJ(2008)Sexual systems and ecological correlates in an azonal tropical forests,SW China.Biotropica,40,160-167.
Darwin C(1877)The Different Forms of Flowers on Plants of the Same Species.John Murray,London.
Diggle PJ(1983)Statistical Analysis of Spatial Point Patterns.Academic Press,New York.
Du RQ(2009)Biostatistics.Higher Education Press,Beijing.(in Chinese)[杜荣骞(2009)生物统计学.高等教育出版社,北京.]
Graff P,Rositano F,Aguiar MR(2013)Changes in sex ratios of a dioecious grass with grazing intensity:The interplay between gender traits,neighbour interactions and spatial patterns.Journal of Ecology,101,1146-1157.
Heikrujam M,Sharma K,Prasad M,Agrawal V(2015)Review on different mechanisms of sex determination and sexlinked molecular markers in dioecious crops-A current update.Euphytica,201,161-194.
Henry IM,Akagi T,Tao R,Comai L(2018)One hundred ways to invent the sexes:Theoretical and observed paths to dioecy in plants.Annual Review of Plant Biology,69,553-575.
Huang SQ,Guo YH(2000)Progress in pollination biology.Chinese Science Bulletin,45,225-237.(in Chinese with English abstract)[黄双全,郭友好(2000)传粉生物学的研究进展.科学通报,45,225-237.]
Hultine KR,Grady KC,Wood TE,Shuster SM,Stella JC,Whitham TG(2016)Climate change perils for dioecious plant species.Nature Plants,2,16109.
James DM(1991)Botany:An Introduction to Plant Biology.Saunders College Publishing,Austin.
Jiang YX,Lu JP(1991)Ecosystem of Tropical Forest of Jianfengling Mountain,Hainan Island,China.Science Press,Beijing.(in Chinese)[蒋有绪,卢俊培(1991)中国海南岛尖峰岭热带林生态系统.科学出版社,北京.]
Jing SW,Coley PD(1990)Dioecy and herbivory:The effect of growth rate on plant defense in Acer negundo.Oikos,58,369-377.
Juvany M,Munné-Bosch S(2015)Sex-related differences in stress tolerance in dioecious plants:A critical appraisal in a physiological context.Journal of Experimental Botany,66,6083-6092.
Li TH,Jiang J,Chen JM,Fan SB(2004)The sexual polymorphism of seed plant.Journal of Northeast Forestry University,32,48-52.(in Chinese with English abstract)[李同华,姜静,陈建名,范士波(2004)种子植物性别的多态性.东北林业大学学报,32,48-52.]
Li YD,Chen BF,Zhou GY,Wu ZM,Zeng QB,Luo TS,Huang SN,Xie MD,Huang Q(2002)Research and Conservation of Tropical Forest and the Biodiversity:A Special Reference to Hainan Island,China.China Forestry Publishing House,Beijing.(in Chinese)[李意德,陈步峰,周光益,吴仲民,曾庆波,骆土寿,黄世能,谢明东,黄全(2002)中国海南岛热带森林及其生物多样性保护研究.中国林业出版社,北京.]
Li YD,Xu H,Luo TS,Chen DX,Lin MX(2012)Bio-Checklist of Jianfengling Station.China Agricultural Press,Beijing.(in Chinese)[李意德,许涵,骆土寿,陈德祥,林明献(2012)尖峰岭生态站生物物种数据集.中国农业出版社,北京.]
Long R(2011)Study on Community Structure of Deciduous Broad-Leaf Forests and Their Plant Sexual System Diversity in Dalaoling,Hubei Province.PhD Dissertation,Beijing Forestry University,Beijing.(in Chinese with English abstract)[龙茹(2011)湖北大老岭落叶阔叶林群落结构和植物性系统多样性研究.博士学位论文,北京林业大学,北京.]
Long R,Shang C,Qu S,Zhang ZX(2011)Distribution pattern of plant sexual system diversity in Populus lasiocarpa Oliv.community.Journal of Beijing Forestry University,33,34-41.(in Chinese with English abstract)[龙茹,尚策,曲上,张志翔(2011)大叶杨群落植物性系统多样性的分布格局.北京林业大学学报,33,34-41.]
Machado LC,Vlopes A,Sazima M(2006)Plant sexual systems and review of the breeding system studies in the Caatinga,a Brazilian tropical dry forest.Annals of Botany,97,277-287.
Maxime RM,Pierre-Olivier C(2015)High incidence of dioecy in young successional tropical forests.Journal of Ecology,103,725-732.
McIntosh RP(1967)The continuum concept of vegetation.The Botanical Review,33,130-187.
Pannell JR(2017)Plant sex determination.Current Biology,27,191-197.
Pielou EC(1975)Ecological Diversity.John Wiley&Sons,New York.
Ramirez N(2005)Plant sexual systems,dichogamy,and herkogamy in the Venezuelan Central Plain.Flora Jena,200,30-48.
Ramirez N,Brito Y(1990)Reproductive biology of a tropical palm swamp community in the Venezuelan LIanos.American Journal of Botany,77,1260-1271.
Renner(2014)The relative and absolute frequencies of angiosperm sexual systems:Dioecy,monoecy,gynodioecy,and an updated online database.American Journal of Botany,101,1588-1596.
Robinson J(2008)The Evolution of Flower Size and Flowering Behaviour in Plants:The Role of Pollination and Pre-dispersal Seed Predation.Master dissertation,University of Southampton,UK.
Shannon CE(1949)Communication theory of secrecy systems.The Bell System Technical Journal,28,656-715.
Simpson EH(1949)Measurement of diversity.Nature,163,688.
Tao J,Zang RG,Li YD,Mao PL,Lin MX(2011)Characteristic of plant functional groups based on ecophysiological traits in a tropical montane rain forest of Hainan Island,South China.Scientia Silvae Sinicae,47(8),14-18.(in Chinese with English abstract)[陶晶,臧润国,李意德,毛培利,林明献(2011)海南尖峰岭热带山地雨林生理生态功能群特征.林业科学,47(8),14-18.]
Xu H,Li YD,Lin MX,Wu JH,Luo TS,Zhou Z,Chen DX,Yang H,Li GJ,Liu SR(2015a)Community characteristics of a 60 ha dynamics plot in the tropical montane rain forest in Jianfengling,Hainan Island.Biodiversity Science,23,192-201.(in Chinese with English abstract)[许涵,李意德,林明献,吴建辉,骆土寿,周璋,陈德祥,杨怀,李广建,刘世荣(2015a)海南尖峰岭热带山地雨林60 ha动态监测样地群落结构特征.生物多样性,23,192-201.]
Xu H,Li YD,Luo TS,Chen DX,Lin MX,Wu JH,Li YP,Yang H,Zhou Z(2015b)Jianfengling Tropical Mountain Rain Forest Dynamic Plot:Community Characteristics,Tree Species and Their Distribution Patterns.China Forestry Publishing House,Beijing.(in Chinese)[许涵,李意德,骆土寿,陈德祥,林明献,吴建辉,李艳朋,杨怀,周璋(2015b)海南尖峰岭热带山地雨林--群落特征、树种及其分布格局.中国林业出版社,北京.]
Xu H,Li YD,Luo TS,Lin MX,Chen DX,Mo JH,Luo W,Huang H(2008)Influence of typhoon Damrey on the tropical montane rain forest community in Jianfengling,Hainan Island,China.Chinese Journal of Plant Ecology,32,1323-1334.(in Chinese with English abstract)[许涵,李意德,骆土寿,林明献,陈德祥,莫锦华,罗文,黄豪(2008)达维台风对海南尖峰岭热带山地雨林群落的影响.植物生态学报,32,1323-1334.]
Xu X,Yang F,Yin CY,Li CY(2007)Research advances in sex-specific responses of dioecious plants to environmental stresses.Chinese Journal of Applied Ecology,18,2626-2631.(in Chinese with English abstract)[胥晓,杨帆,尹春英,李春阳(2007)雌雄异株植物对环境胁迫响应的性别差异研究进展.应用生态学报,18,2626-2631.]
Yamakura T,Kanzaki M,Itoh A,Ohkubo T,Ogino K,Chai EOK,Lee HS,Ashton PS(1995)Topography of a large-scale research plot established within a tropical rain forest at Lambir,Sarawak.Tropics,5,41-56.
Yan LH(2007)Sexual system and environmental adaptability of vines in Hupingshan Mountain,Hunan Province.Journal of Northeast Forestry University,35,35-36,39.(in Chinese with English abstract)[颜立红(2007)湖南壶瓶山藤本植物的有性系统及其环境适应性.东北林业大学学报,35,35-36,39.]
Yin CY,Li CY(2007)Gender differences of dioecious plants related sex ratio:Recent advances and future prospects.Chinese Journal of Applied Environmental Biology,13,419-425.(in Chinese with English abstract)[尹春英,李春阳(2007)雌雄异株植物与性别比例有关的性别差异研究现状与展望.应用与环境生物学报,13,419-425.]
Zeng QB,Li YD,Chen BF,Wu ZM,Zhou GY(1997)Research and Management of Tropical Forest Ecosystem.China Forestry Publishing House,Beijing.(in Chinese)[曾庆波,李意德,陈步峰,吴仲民,周光益(1997)热带森林生态系统研究与管理.中国林业出版社,北京.]
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