海岛植物不同演替阶段植物功能性状与环境因子的变化规律
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摘要
植物功能性状与环境之间的关系是功能性状研究的重点,环境因子驱使植物功能性状发生变化,进而推动群落发生演替。以平潭岛4个不同演替阶段的森林植被(灌草丛、针叶林、针阔混交林、常绿阔叶林)为研究对象,结合不同群落演替阶段的物种特征和群落结构,分析海岛不同演替阶段茎、叶功能性状的变化规律,以及功能性状与环境因子的关系。结果表明:(1)随着演替的进行,土壤养分和水分逐渐增加,土壤pH逐渐下降。比叶面积(SLA)、叶片氮含量(LNC)、叶片磷含量(LPC)、茎氮含量(SNC)、茎磷含量(SPC)下降后上升,叶厚度(LT)、叶片碳含量(LCC)、茎碳含量(SCC)与之相反,叶干物质含量(LDMC)、茎组织密度(STD)逐渐上升。(2)冗余分析表明,演替早期植物主要分布在土壤pH、容重高的贫瘠环境,拥有较高SLA、SNC、SPC、LPC的性状组合;演替后期植物主要分布在土壤养分和水分高的肥沃环境,拥有较高的STD、LDMC、LCC、LNC的性状组合。其中,土壤有机质和全氮含量是影响海岛植物演替过程中功能性状变化的关键环境因子。研究海岛植物功能性状与环境之间的关系随演替的变化规律,探讨各演替阶段功能性状和环境特征,以及功能性状如何响应环境变化。旨在为今后选择合适的树种进行海岛植被修复和重建提供依据。
The relationship between plant functional traits and environment is the focus of functional traits research. Environmental factors drive the changes in plant functional traits, which in turn promote community succession. Based on the forest vegetation of four different succession stages(shrub-grassland, coniferous forest, mixed wood, and broadleaf forest) in Pingtan Island, this study combined the species characteristics and community structure of different community successional stages. We analyzed the changes in stem and leaf functional traits in different succession stages of island plants. Moreover, we explored the relationship between plant functional traits and environmental factors. The results showed the following.(1) With progress in succession, soil nutrient and water content gradually increased and the soil pH gradually decreased. The specific leaf area(SLA), leaf nitrogen content(LNC), leaf phosphorus content(LPC), stem nitrogen content(SNC), and stem phosphorus content(SPC) decreased initially, and then increased, Meanwhile, the leaf thickness(LT), leaf carbon content(LCC), and stem carbon content(SCC) increased initially, and then decreased. The leaf dry matter content(LDMC) and stem tissue density(STD) gradually increased.(2) The redundancy analysis showed that the early succession plants were mainly distributed in barren environments with high soil pH and high bulk density, and they presented higher SLA, SNC, SPC, and LPC. Plants in the late succession stages were mainly distributed in fertile environments with high soil nutrient and water content, and they presented higher STD, LDMC, LCC, and LNC. Therefore, soil organic matter and total nitrogen are the important environmental factors that affect the functional traits of island plants succession. We studied the relationship between plant functional traits and environment along with the changes in succession, understood the functional traits and environmental characteristics of each succession stage, and revealed how plant functional traits adapted to the environmental changes. The study provides the basis for the selection of suitable tree species for island vegetation restoration and reconstruction in the future.
The relationship between plant functional traits and environment is the focus of functional traits research. Environmental factors drive the changes in plant functional traits, which in turn promote community succession. Based on the forest vegetation of four different succession stages(shrub-grassland, coniferous forest, mixed wood, and broadleaf forest) in Pingtan Island, this study combined the species characteristics and community structure of different community successional stages. We analyzed the changes in stem and leaf functional traits in different succession stages of island plants. Moreover, we explored the relationship between plant functional traits and environmental factors. The results showed the following.(1) With progress in succession, soil nutrient and water content gradually increased and the soil pH gradually decreased. The specific leaf area(SLA), leaf nitrogen content(LNC), leaf phosphorus content(LPC), stem nitrogen content(SNC), and stem phosphorus content(SPC) decreased initially, and then increased, Meanwhile, the leaf thickness(LT), leaf carbon content(LCC), and stem carbon content(SCC) increased initially, and then decreased. The leaf dry matter content(LDMC) and stem tissue density(STD) gradually increased.(2) The redundancy analysis showed that the early succession plants were mainly distributed in barren environments with high soil pH and high bulk density, and they presented higher SLA, SNC, SPC, and LPC. Plants in the late succession stages were mainly distributed in fertile environments with high soil nutrient and water content, and they presented higher STD, LDMC, LCC, and LNC. Therefore, soil organic matter and total nitrogen are the important environmental factors that affect the functional traits of island plants succession. We studied the relationship between plant functional traits and environment along with the changes in succession, understood the functional traits and environmental characteristics of each succession stage, and revealed how plant functional traits adapted to the environmental changes. The study provides the basis for the selection of suitable tree species for island vegetation restoration and reconstruction in the future.
引文
[1] Violle C,Navas M L,Vile D,Kazakou E,Fortunel C,Hummel I,Garnier E.Let the concept of trait be functional.Oikos,2007,116(5):882- 892.
[2] 孟婷婷,倪健,王国宏.植物功能性状与环境和生态系统功能.植物生态学报,2007,31(1):150- 165.
[3] 胡耀升,么旭阳,刘艳红.长白山不同演替阶段森林植物功能性状及其与地形因子间的关系.生态学报,2014,34(20):5915- 5924.
[4] Wright I J,Reich P B,Westoby M,Ackerly D D,Baruch Z,Bongers F,Cavender-Bares J,Chapin T,Cornelissen J H C,Diemer M,Flexas J,Garnier E,Groom P K,Gulias J,Hikosaka K,Lamont B B,Lee T,Lee W,Lusk C,Midgley J J,Navas M L,Niinemets ü,Oleksyn J,Osada N,Poorter H,Poot P,Prior L,Pyankov V I,Roumet C,Thomas S C,Tjoelker M G,Veneklaas E J,Villar R.The worldwide leaf economics spectrum.Nature,2004,428(6985):821- 827.
[5] 路兴慧,丁易,臧润国,邹正冲,黄卢标.海南岛热带低地雨林老龄林木本植物幼苗的功能性状分析.植物生态学报,2011,35(12):1300- 1309.
[6] 李丹,康萨如拉,赵梦颖,张庆,任海娟,任婧,周俊梅,王珍,吴仁吉,牛建明.内蒙古羊草草原不同退化阶段土壤养分与植物功能性状的关系.植物生态学报,2016,40(10):991- 1002.
[7] 刘贵峰,刘玉平,达福白乙拉,程伟燕,高学磊,姜丽丽.大青沟自然保护区主要森林群落优势种的叶性状.生态学报,2017,37(14):4646- 4655.
[8] 盘远方,陈兴彬,姜勇,梁士楚,陆志任,黄宇欣,倪鸣源,覃彩丽,刘润红.桂林岩溶石山灌丛植物叶功能性状和土壤因子对坡向的响应.生态学报,2018,38(5):1581- 1589.
[9] Chai Y F,Yue M,Wang M,Xu J S,Liu X,Zhang R C,Wan P C.Plant functional traits suggest a change in novel ecological strategies for dominant species in the stages of forest succession.Oecologia,2016,180(3):771- 783.
[10] Kahmen S,Poschlod P.Plant functional trait responses to grassland succession over 25 Years.Journal of Vegetation Science,2010,15(1):21- 32.
[11] 许洺山,黄海侠,史青茹,杨晓东,周刘丽,赵延涛,张晴晴,阎恩荣.浙东常绿阔叶林植物功能性状对土壤含水量变化的响应.植物生态学报,2015,39(9):857- 866.
[12] Muscarella R,Uriarte M,Aide T M,Erickson D L.Functional convergence and phylogenetic divergence during secondary succession of subtropical wet forests in Puerto Rico.Journal of Vegetation Science,2016,27(2):283- 294.
[13] Castro H,Lehsten V,Lavorel S,Freitas H.Functional response traits in relation to land use change in the Montado.Agriculture,Ecosystems & Environment,2010,137(1/2):183- 191.
[14] Reich P B,Wright I J,Cavender-Bares J,Craine J M,Oleksyn J,Westoby M,Walters M B.The evolution of plant functional variation:traits,spectra,and strategies.International Journal of Plant Sciences,2003,164(S3):S143-S164.
[15] Craven D,Hall J S,Berlyn G P,Ashton M,Van Breugel M.Changing gears during succession:shifting functional strategies in young tropical secondary forests.Oecologia,2015,179(1):293- 305.
[16] Stuessy T F,Jakubowsky G,Gómez R S,Pfosser M,Schlüter P M,Fer T,Sun B Y,Kato H.Anagenetic evolution in island plants.Journal of Biogeography,2010,33(7):1259- 1265.
[17] 黄柳菁,张增可,郑俊鸣,黄少君,林丽丽,王齐,邓传远,刘兴诏.大陆性海岛野生植物功能性状特征及其演变趋势——以平潭岛为例.热带亚热带植物学报,2017,25(5):429- 437.
[18] Poorter L.Growth responses of 15 rain-forest tree species to a light gradient:the relative importance of morphological and physiological traits.Functional Ecology,1999,13(3):396- 410.
[19] Méndez-Alonzo R,Ewers F W,Sack L.Ecological variation in leaf biomechanics and its scaling with tissue structure across three mediterranean‐climate plant communities.Functional Ecology,2013,27(2):544- 554.
[20] Cornelissen J H C,Lavorel S,Garnier E,Díaz S,Buchmann N,Gurvich D E,Reich P B,Ter Steege H,Morgan H D,Van Der Heijden M G A,Pausas J G,Poorter H.A handbook of protocols for standardised and easy measurement of plant functional traits worldwide.Australian Journal of Botany,2003,51(4):335- 380.
[21] 卜文圣,臧润国,丁易,张俊艳,阮云泽.海南岛热带低地雨林群落水平植物功能性状与环境因子相关性随演替阶段的变化.生物多样性,2013,21(3):278- 287.
[22] Lep? J,?milauer P.Multivariate Analysis of Ecological Data Using CANOCO.Cambridge:Cambridge University Press,2003:1149-166.
[23] 黄永涛.亚热带常绿落叶阔叶混交林的功能性群落结构及其动态研究[D].北京:中国林业科学研究院,2016.
[24] 赵文霞.亚热带常绿阔叶林常见树种根茎叶功能性状研究[D].北京:北京林业大学,2016.
[25] 丁佳,吴茜,闫慧,张守仁.地形和土壤特性对亚热带常绿阔叶林内植物功能性状的影响.生物多样性,2011,19(2):158- 167.
[26] Westoby M,Falster D S,Moles A T,Vesk P A,Wright I J.Plant ecological strategies:some leading dimensions of variation between species.Annual Review of Ecology and Systematics,2002,33(1):125- 159.
[27] Reich P B,Walters M B,Ellsworth D S.From tropics to tundra:global convergence in plant functioning.Proceedings of the National Academy of Sciences of the United States of America,1997,94(25):13730- 13734.
[28] 张俊艳.海南岛热带天然针叶林—阔叶林交错区的群落特征研究[D].北京:中国林业科学研究院,2014.
[29] Vitousek P M,Porder S,Houlton B Z,Chadwick O A.Terrestrial phosphorus limitation:mechanisms,implications,and nitrogen-phosphorus interactions.Ecological Applications,2010,20(1):5- 15.
[30] 韩文轩,吴漪,汤璐瑛,陈雅涵,李利平,贺金生,方精云.北京及周边地区植物叶的碳氮磷元素计量特征.北京大学学报:自然科学版,2008,45(4):855- 860.
[31] 宋光满,韩涛涛,洪岚,张玲玲,李晓波,任海.演替过程中植物功能性状研究进展.生态科学,2018,37(2):207- 213.
[32] Zhang J Y,Cheng K W,Zang R G,Ding Y.Environmental filtering of species with different functional traits into plant assemblages across a tropical coniferous-broadleaved forest ecotone.Plant and Soil,2014,380(1/2):361- 374.
[33] Ackerly D,Knight C,Weiss S,Barton K,Starmer K.Leaf size,specific leaf area and microhabitat distribution of chaparral woody plants:contrasting patterns in species level and community level analyses.Oecologia,2002,130(3):449- 457.
[34] Franklin J,Spears-Lebrun L A,Deutschman D H,Marsden K.Impact of a high-intensity fire on mixed evergreen and mixed conifer forests in the Peninsular Ranges of southern California,USA.Forest Ecology and Management,2006,235(1/3):18- 29.
[2] 孟婷婷,倪健,王国宏.植物功能性状与环境和生态系统功能.植物生态学报,2007,31(1):150- 165.
[3] 胡耀升,么旭阳,刘艳红.长白山不同演替阶段森林植物功能性状及其与地形因子间的关系.生态学报,2014,34(20):5915- 5924.
[4] Wright I J,Reich P B,Westoby M,Ackerly D D,Baruch Z,Bongers F,Cavender-Bares J,Chapin T,Cornelissen J H C,Diemer M,Flexas J,Garnier E,Groom P K,Gulias J,Hikosaka K,Lamont B B,Lee T,Lee W,Lusk C,Midgley J J,Navas M L,Niinemets ü,Oleksyn J,Osada N,Poorter H,Poot P,Prior L,Pyankov V I,Roumet C,Thomas S C,Tjoelker M G,Veneklaas E J,Villar R.The worldwide leaf economics spectrum.Nature,2004,428(6985):821- 827.
[5] 路兴慧,丁易,臧润国,邹正冲,黄卢标.海南岛热带低地雨林老龄林木本植物幼苗的功能性状分析.植物生态学报,2011,35(12):1300- 1309.
[6] 李丹,康萨如拉,赵梦颖,张庆,任海娟,任婧,周俊梅,王珍,吴仁吉,牛建明.内蒙古羊草草原不同退化阶段土壤养分与植物功能性状的关系.植物生态学报,2016,40(10):991- 1002.
[7] 刘贵峰,刘玉平,达福白乙拉,程伟燕,高学磊,姜丽丽.大青沟自然保护区主要森林群落优势种的叶性状.生态学报,2017,37(14):4646- 4655.
[8] 盘远方,陈兴彬,姜勇,梁士楚,陆志任,黄宇欣,倪鸣源,覃彩丽,刘润红.桂林岩溶石山灌丛植物叶功能性状和土壤因子对坡向的响应.生态学报,2018,38(5):1581- 1589.
[9] Chai Y F,Yue M,Wang M,Xu J S,Liu X,Zhang R C,Wan P C.Plant functional traits suggest a change in novel ecological strategies for dominant species in the stages of forest succession.Oecologia,2016,180(3):771- 783.
[10] Kahmen S,Poschlod P.Plant functional trait responses to grassland succession over 25 Years.Journal of Vegetation Science,2010,15(1):21- 32.
[11] 许洺山,黄海侠,史青茹,杨晓东,周刘丽,赵延涛,张晴晴,阎恩荣.浙东常绿阔叶林植物功能性状对土壤含水量变化的响应.植物生态学报,2015,39(9):857- 866.
[12] Muscarella R,Uriarte M,Aide T M,Erickson D L.Functional convergence and phylogenetic divergence during secondary succession of subtropical wet forests in Puerto Rico.Journal of Vegetation Science,2016,27(2):283- 294.
[13] Castro H,Lehsten V,Lavorel S,Freitas H.Functional response traits in relation to land use change in the Montado.Agriculture,Ecosystems & Environment,2010,137(1/2):183- 191.
[14] Reich P B,Wright I J,Cavender-Bares J,Craine J M,Oleksyn J,Westoby M,Walters M B.The evolution of plant functional variation:traits,spectra,and strategies.International Journal of Plant Sciences,2003,164(S3):S143-S164.
[15] Craven D,Hall J S,Berlyn G P,Ashton M,Van Breugel M.Changing gears during succession:shifting functional strategies in young tropical secondary forests.Oecologia,2015,179(1):293- 305.
[16] Stuessy T F,Jakubowsky G,Gómez R S,Pfosser M,Schlüter P M,Fer T,Sun B Y,Kato H.Anagenetic evolution in island plants.Journal of Biogeography,2010,33(7):1259- 1265.
[17] 黄柳菁,张增可,郑俊鸣,黄少君,林丽丽,王齐,邓传远,刘兴诏.大陆性海岛野生植物功能性状特征及其演变趋势——以平潭岛为例.热带亚热带植物学报,2017,25(5):429- 437.
[18] Poorter L.Growth responses of 15 rain-forest tree species to a light gradient:the relative importance of morphological and physiological traits.Functional Ecology,1999,13(3):396- 410.
[19] Méndez-Alonzo R,Ewers F W,Sack L.Ecological variation in leaf biomechanics and its scaling with tissue structure across three mediterranean‐climate plant communities.Functional Ecology,2013,27(2):544- 554.
[20] Cornelissen J H C,Lavorel S,Garnier E,Díaz S,Buchmann N,Gurvich D E,Reich P B,Ter Steege H,Morgan H D,Van Der Heijden M G A,Pausas J G,Poorter H.A handbook of protocols for standardised and easy measurement of plant functional traits worldwide.Australian Journal of Botany,2003,51(4):335- 380.
[21] 卜文圣,臧润国,丁易,张俊艳,阮云泽.海南岛热带低地雨林群落水平植物功能性状与环境因子相关性随演替阶段的变化.生物多样性,2013,21(3):278- 287.
[22] Lep? J,?milauer P.Multivariate Analysis of Ecological Data Using CANOCO.Cambridge:Cambridge University Press,2003:1149-166.
[23] 黄永涛.亚热带常绿落叶阔叶混交林的功能性群落结构及其动态研究[D].北京:中国林业科学研究院,2016.
[24] 赵文霞.亚热带常绿阔叶林常见树种根茎叶功能性状研究[D].北京:北京林业大学,2016.
[25] 丁佳,吴茜,闫慧,张守仁.地形和土壤特性对亚热带常绿阔叶林内植物功能性状的影响.生物多样性,2011,19(2):158- 167.
[26] Westoby M,Falster D S,Moles A T,Vesk P A,Wright I J.Plant ecological strategies:some leading dimensions of variation between species.Annual Review of Ecology and Systematics,2002,33(1):125- 159.
[27] Reich P B,Walters M B,Ellsworth D S.From tropics to tundra:global convergence in plant functioning.Proceedings of the National Academy of Sciences of the United States of America,1997,94(25):13730- 13734.
[28] 张俊艳.海南岛热带天然针叶林—阔叶林交错区的群落特征研究[D].北京:中国林业科学研究院,2014.
[29] Vitousek P M,Porder S,Houlton B Z,Chadwick O A.Terrestrial phosphorus limitation:mechanisms,implications,and nitrogen-phosphorus interactions.Ecological Applications,2010,20(1):5- 15.
[30] 韩文轩,吴漪,汤璐瑛,陈雅涵,李利平,贺金生,方精云.北京及周边地区植物叶的碳氮磷元素计量特征.北京大学学报:自然科学版,2008,45(4):855- 860.
[31] 宋光满,韩涛涛,洪岚,张玲玲,李晓波,任海.演替过程中植物功能性状研究进展.生态科学,2018,37(2):207- 213.
[32] Zhang J Y,Cheng K W,Zang R G,Ding Y.Environmental filtering of species with different functional traits into plant assemblages across a tropical coniferous-broadleaved forest ecotone.Plant and Soil,2014,380(1/2):361- 374.
[33] Ackerly D,Knight C,Weiss S,Barton K,Starmer K.Leaf size,specific leaf area and microhabitat distribution of chaparral woody plants:contrasting patterns in species level and community level analyses.Oecologia,2002,130(3):449- 457.
[34] Franklin J,Spears-Lebrun L A,Deutschman D H,Marsden K.Impact of a high-intensity fire on mixed evergreen and mixed conifer forests in the Peninsular Ranges of southern California,USA.Forest Ecology and Management,2006,235(1/3):18- 29.