同倍体杂交物种紫果云杉的生态位分化及其未来潜在分布区预测
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摘要
【目的】以我国特有的同倍体杂交树种紫果云杉为对象,研究其与亲本树种丽江云杉和青杄间环境生态位的差异,并预测其在未来气候变化下21世纪50年代(2050s)和80年代(2080s)潜在分布区的变化。【方法】在收集紫果云杉及其亲本种地理分布信息的基础上,利用地理信息系统(Arc GIS)技术获取相应生境的环境变量,通过Kruskal-Wallis多重秩和检验、判别式分析(DFA)和主成分分析(PCA)等统计方法量化分析云杉属3个树种的生态位差异,并利用最大熵模型(Max Ent)结合3种大气环流模型(BCC-CSM1-1、CCCma_Can ESM2和CSIROMk3.6.0)模拟3个树种在3种气候变化情景(即温室气体最低排放,RCP2.6;中度稳定排放,RCP4.5;高度排放,RCP 8.5)下未来2050s和2080s的潜在分布。【结果】Kruskal-Wallis检验、DFA和PCA结果均表明紫果云杉生境的水热条件与亲本种间均存在显著差异。对水分条件而言,尽管紫果云杉生境的最冷季与最暖季降雨量居于亲本种之间,但其生境土壤湿度显著高于2个亲本种;对热量条件而言,紫果云杉生境的最冷月最低温显著低于2个亲本种,且地面结霜频率显著高于亲本。进一步对云杉属3个树种的未来潜在分布区模拟显示,紫果云杉仅在RCP2.6下2080s的潜在分布面积与当前相比略有缩减(约5%),而在此情景下2050s的潜在分布区面积和其余2个情景下2050s和2080s的潜在分布面积高于当前分布区面积。综合分析所有情景及时间段后发现,紫果云杉未来潜在分布区面积平均增加17%以上,且分布区明显呈现由青藏高原东南部边缘向内部扩张的趋势。而其亲本种青杄受气候变暖威胁严重,其潜在分布面积(综合所有情景及时间段)平均减少了21%以上,丽江云杉除在RCP4.5和RCP8.5下2080s的潜在分布面积稍高于当前(不到2%)外,其余情景下的潜在分布面积也都低于当前,其潜在分布面积平均减少5%左右(综合所有情景及时间段)。【结论】明确了同倍体杂交物种紫果云杉与其亲本种生态位的环境差异,即高土壤湿度和冬季低温是紫果云杉与其亲本种产生生态位分化的主要因子;且在未来温度显著增加背景下,基于MaxEnt模型预测表明紫果云杉在未来2050s和2080s潜在分布区面积将显著增加,推测其在未来将发挥更重要的生态安全屏障作用。
【Objective】In this study,we analyzed the environmental divergence between the endemic homoploid hybrid tree species Picea purpurea and its progenitors P. wilsonii and P. likiangensis and predicted the change of their potential distribution under climate change for two future periods: 2050 s and 2080 s. 【Method 】 On the basis of collected geographical distributional information,the geographical information system software( Arc GIS) was used to obtain the environmental variables which was related to the distribution of P. purpurea and its parental species. Then the ecological divergence was quantified via Kruskal-Wallis multiple-range test,discriminant function analysis( DFA),and principle components analysis( PCA) etc. The distribution patterns of three Picea species under different climate change scenarios( i. e. the lowest,moderate,and the highest greenhouse gas emission scenario; RCP2. 6,RCP4. 5 and RCP8. 5) were estimated using maximum entropy analysis( Max Ent) and three general circulation models( BCC-CSM1-1,CCCma_Can ESM2 和 CSIRO-Mk3. 6. 0) for the 2050 s and the 2080 s.【Result】 The results from Kruskal-Wallis tests,DFA and PCA indicated that the water and heat availability for P. purpurea were all significantly different from its parental species.That is, despite of the precipitation of coldest and warmest quarter of P. purpurea was intermediate between its progenitors,its soil moisture was significantly higher than them; the min temperature of coldest month of P. purpurea was significantly lower than that of its parental species,and the ground-frost frequency of P. purpurea was inversely higher than that of its progenitors. Furthermore,results from simulation-based estimates revealed that the potential distribution area for P. purpurea would decrease( about 5%) in the 2080 s only under RCP2. 6,while in the 2050 s of this scenario and under other two scenarios,its distributional area in the two time periods were higher than the current. After integrated all three scenarios and two time periods,the potential distribution area of P. purpurea would increase by an average of17% in the future,and exhibited an expansion tendency from the edge of southeast of Tibetan Plateau to its northeast inner. In contrast,the potential distribution of P. wilsonii would be severely threated by the climate change,its distributional area would decrease by average of more than 21% in the future( after integrated all three scenarios and two time periods). The potential distributional area of P. likiangensis would decrease by an average of 5% for all scenarios and periods in the future than current( after integrated all three scenarios and two time periods),excepted for the 2080 s under both RCP4. 5 and 8. 5( slightly higher less than 2%). 【Conclusion 】 We demonstrated the environmental divergence between the homoploid hybrid species P. purpurea and its parental species in this study: the higher soil moisture and the low temperature in the winter were the key factors that driving the ecological divergence in P. purpurea from its progenitors. The potential distribution area of P. purpurea would significantly increase in the 2050 s and 2080 s.This suggests that P. purpurea plays an important role in ecological security and protection in the future.
【Objective】In this study,we analyzed the environmental divergence between the endemic homoploid hybrid tree species Picea purpurea and its progenitors P. wilsonii and P. likiangensis and predicted the change of their potential distribution under climate change for two future periods: 2050 s and 2080 s. 【Method 】 On the basis of collected geographical distributional information,the geographical information system software( Arc GIS) was used to obtain the environmental variables which was related to the distribution of P. purpurea and its parental species. Then the ecological divergence was quantified via Kruskal-Wallis multiple-range test,discriminant function analysis( DFA),and principle components analysis( PCA) etc. The distribution patterns of three Picea species under different climate change scenarios( i. e. the lowest,moderate,and the highest greenhouse gas emission scenario; RCP2. 6,RCP4. 5 and RCP8. 5) were estimated using maximum entropy analysis( Max Ent) and three general circulation models( BCC-CSM1-1,CCCma_Can ESM2 和 CSIRO-Mk3. 6. 0) for the 2050 s and the 2080 s.【Result】 The results from Kruskal-Wallis tests,DFA and PCA indicated that the water and heat availability for P. purpurea were all significantly different from its parental species.That is, despite of the precipitation of coldest and warmest quarter of P. purpurea was intermediate between its progenitors,its soil moisture was significantly higher than them; the min temperature of coldest month of P. purpurea was significantly lower than that of its parental species,and the ground-frost frequency of P. purpurea was inversely higher than that of its progenitors. Furthermore,results from simulation-based estimates revealed that the potential distribution area for P. purpurea would decrease( about 5%) in the 2080 s only under RCP2. 6,while in the 2050 s of this scenario and under other two scenarios,its distributional area in the two time periods were higher than the current. After integrated all three scenarios and two time periods,the potential distribution area of P. purpurea would increase by an average of17% in the future,and exhibited an expansion tendency from the edge of southeast of Tibetan Plateau to its northeast inner. In contrast,the potential distribution of P. wilsonii would be severely threated by the climate change,its distributional area would decrease by average of more than 21% in the future( after integrated all three scenarios and two time periods). The potential distributional area of P. likiangensis would decrease by an average of 5% for all scenarios and periods in the future than current( after integrated all three scenarios and two time periods),excepted for the 2080 s under both RCP4. 5 and 8. 5( slightly higher less than 2%). 【Conclusion 】 We demonstrated the environmental divergence between the homoploid hybrid species P. purpurea and its parental species in this study: the higher soil moisture and the low temperature in the winter were the key factors that driving the ecological divergence in P. purpurea from its progenitors. The potential distribution area of P. purpurea would significantly increase in the 2050 s and 2080 s.This suggests that P. purpurea plays an important role in ecological security and protection in the future.
引文
梁玉莲,延晓冬.2016.RCPs情景下中国21世纪气候变化预估及不确定性分析.热带气象学报,32(2):183-192.(Liang Y L,Yan X D.2016.Prediction of climate change over China and uncertainty analysis during the 21stcentury under RCPs.Journal of Tropical Meteorology,32(2):183-192.[in Chinese])
孙永帅.2012.紫果云杉的二倍体杂交起源.兰州:兰州大学博士学位论文.(Sun Y S.2012.Diploid hybrid origin of Picea purpurea.Lanzhou:Ph Dthesis of Lanzhou University.[in Chinese])
殷晓洁,周广胜,隋兴华,等.2013.辽东栎林潜在地理分布及其主导因子.林业科学,49(8):10-14.(Yin Y J,Zhou G S,Sui X H,et al.2013.Potential geographical distribution of Quercus wutaishanica forest and its dominant factors.Scientia Silvae Science,49(8):10-14.[in Chinese])
张雷,刘世荣,孙鹏森,等.2011.气候变化对物种分布影响模拟中的不确定性组分分割与制图---以油松为例.生态学报,31(19):5749-5761.(Zhang L,Liu S R,Sun P S,et al.2011.Partitioning and mapping the sources of variations in the ensemble forecasting of species distribution under climate change:a case study of Pinus tabulaeformis.Acta Ecologica Sinica,31(19):5749-5761.[in Chinese])
郑万钧.1938.中国树木志:第一卷.北京:中国林业出版社,221-223.(Zheng W J.1938.Chinese tree annals:Volume I.Beijing:China Forestry Publishing House,221-223.[in Chinese])
中国森林编辑委员会.1998.中国森林:第二卷.北京:中国林业出版社,749-750.(China Forest Editorial Board.1998.China forest:VolumeⅡ.Beijing:China Forestry Publishing House,749-750.[in Chinese])
Forster P,Ramaswamy V,Artaxo P,et al.2007.Changes in atmospheric constituents and in radiative forcing∥Climate change 2007:The physical science basis.Contribution of Working Group I to the 4th Assessment Report of the Intergovernmental Panel on Climate Change,DLR:129-234.
Garzón M B,Dios R S,Ollero H S.2008.Effects of climate change on the distribution of Iberian tree species.Applied Vegetation Science,11(2):169-178.
Gompert Z,Fordyce J A,Forister M L,et al.2006.Homoploid speciation in an extreme habitat.Science,314(5807):1923-1925.
Gregory M F,Helen M M,Kristen L C,et al.2010.Armillaria root disease-caused tree mortality following silvicultural treatments(shelterwood or group selection)in an Oregon mixed-conifer forest:insights from a 10-year case study.Western Journal of Applied Forestry,25(3):136-143.
Guisan A,Thuiller W.2005.Predicting species distribution:offering more than simple habitat models.Ecology Letters,8(9):993-1009.
Lexer C,Welch M E,Raymond O,et al.2003.The origin of ecological divergence in Helianthus paradoxus(Asteraceae):selection on transgressive characters in a novel hybrid habitat.Evolution,57(9):1989-2000.
Ma F,Zhang X W,Chen L T,et al.2013.The alpine homoploid hybrid Pinus densata has greater cold photosynthesis tolerance than its progenitors.Environmental&Experimental Botany,85:85-91.
Ma F,Zhao C,Milne R,et al.2010.Enhanced drought-tolerance in the homoploid hybrid species Pinus densata:implication for its habitat divergence from two progenitors.New Phytologist,185(1):204-216.
Mallet J.2005.Hybridization as an invasion of the genome.Trends in Ecology&Evolution,20(5):229-237.
Mantgem P J V,Stephenson N L,Byrne J C,et al.2009.Widespread increase of tree mortality rates in the western United States.Science,323(5913):521-524.
Mao J F,Wang X R.2011.Distinct niche divergence characterizes the homoploid hybrid speciation of Pinus densata on the Tibetan plateau.American Naturalist,177(4):424-439.
Meinshausen M,Smith S J,Calvin K,et al.2011.The RCP greenhouse gas concentrations and their extensions from 1765 to 2300.Climatic Change,109(1):213-241.
Pearson R G,Dawson T P.2003.Predicting the impacts of climate change on the distribution of species:are bioclimate envelope models useful?Global Ecology&Biogeography,12(5):361-371.
Phillips S J,Anderson R P,Schapire R E.2008.Modeling of species distributions with Max Ent:new extensions and a comprehensive evaluation.Ecography,31(2):161-175.
R Development Core Team.2014.R:A language and environment for statistical computing.R foundation for statistical computing.Vienna,Austria.
Ren Z P,Wang D Q,Ma A M,et al.2016.Predicting malaria vector distribution under climate change scenarios in China:challenges for malaria elimination.Scientific Reports,6:20604.
Rieseberg L H,Archer M A,Wayne R K.1999.Transgressive segregation,adaptation and speciation.Heredity,83(4):363-372.
Rieseberg L H,Lexer C.2003.Major ecological transitions in wild sunflowers facilitated by hybridization.Science,301(5637):1211-1216.
Rieseberg L H.1997.Hybrid origins of plant species.Annual Review of Ecology and Systematics,28(1):359-389.
Rundle H D,Nosil P.2005.Ecological speciation.Ecology Letters,8(3):336-352.
Schumer M,Rosenthal G G,Andolfatto P.2014.How common is homoploid hybrid speciation?Evolution,68(6):1553-1560.
Schwarzbach A E,Donovan L A,Rieseberg L H.2001.Transgressive character expression in a hybrid sunflower species.American Journal of Botany,88(2):270-277.
Sun Y,Abbott R J,Li L,et al.2014.Evolutionary history of Purple cone spruce(Picea purpurea)in the Qinghai-Tibet Plateau:homoploid hybrid origin and Pleistocene expansion.Molecular Ecology,23(2):343-359.
Szewczyk J.2009.Tree mortality and effects of release from competition in an old-growth Fagus-Abies-Picea stand.Journal of Vegetation Science,12(5):621-626.
Wang B,Mao J,Gao J,et al.2011.Colonization of the Tibetan Plateau by the homoploid hybrid pine Pinus densata.Molecular Ecology,20(18):3796-3811.
Xu D,Yan H.2001.A study of the impacts of climate change on the geographic distribution of Pinus koraiensis in China.Environment International,27(2):201-205.
Xu Y,Xu C H.2012.Preliminary assessment of simulations of climate changes over China by CMIP5 multi-models.Atmospheric and Oceanic Science Letters,5(6):489-494.
Yakimowski S B,Rieseberg L H.2014.The role of homoploid hybridization in evolution:a century of studies synthesizing genetics and ecology.American Journal of Botany,101(8):1247-1258.
孙永帅.2012.紫果云杉的二倍体杂交起源.兰州:兰州大学博士学位论文.(Sun Y S.2012.Diploid hybrid origin of Picea purpurea.Lanzhou:Ph Dthesis of Lanzhou University.[in Chinese])
殷晓洁,周广胜,隋兴华,等.2013.辽东栎林潜在地理分布及其主导因子.林业科学,49(8):10-14.(Yin Y J,Zhou G S,Sui X H,et al.2013.Potential geographical distribution of Quercus wutaishanica forest and its dominant factors.Scientia Silvae Science,49(8):10-14.[in Chinese])
张雷,刘世荣,孙鹏森,等.2011.气候变化对物种分布影响模拟中的不确定性组分分割与制图---以油松为例.生态学报,31(19):5749-5761.(Zhang L,Liu S R,Sun P S,et al.2011.Partitioning and mapping the sources of variations in the ensemble forecasting of species distribution under climate change:a case study of Pinus tabulaeformis.Acta Ecologica Sinica,31(19):5749-5761.[in Chinese])
郑万钧.1938.中国树木志:第一卷.北京:中国林业出版社,221-223.(Zheng W J.1938.Chinese tree annals:Volume I.Beijing:China Forestry Publishing House,221-223.[in Chinese])
中国森林编辑委员会.1998.中国森林:第二卷.北京:中国林业出版社,749-750.(China Forest Editorial Board.1998.China forest:VolumeⅡ.Beijing:China Forestry Publishing House,749-750.[in Chinese])
Forster P,Ramaswamy V,Artaxo P,et al.2007.Changes in atmospheric constituents and in radiative forcing∥Climate change 2007:The physical science basis.Contribution of Working Group I to the 4th Assessment Report of the Intergovernmental Panel on Climate Change,DLR:129-234.
Garzón M B,Dios R S,Ollero H S.2008.Effects of climate change on the distribution of Iberian tree species.Applied Vegetation Science,11(2):169-178.
Gompert Z,Fordyce J A,Forister M L,et al.2006.Homoploid speciation in an extreme habitat.Science,314(5807):1923-1925.
Gregory M F,Helen M M,Kristen L C,et al.2010.Armillaria root disease-caused tree mortality following silvicultural treatments(shelterwood or group selection)in an Oregon mixed-conifer forest:insights from a 10-year case study.Western Journal of Applied Forestry,25(3):136-143.
Guisan A,Thuiller W.2005.Predicting species distribution:offering more than simple habitat models.Ecology Letters,8(9):993-1009.
Lexer C,Welch M E,Raymond O,et al.2003.The origin of ecological divergence in Helianthus paradoxus(Asteraceae):selection on transgressive characters in a novel hybrid habitat.Evolution,57(9):1989-2000.
Ma F,Zhang X W,Chen L T,et al.2013.The alpine homoploid hybrid Pinus densata has greater cold photosynthesis tolerance than its progenitors.Environmental&Experimental Botany,85:85-91.
Ma F,Zhao C,Milne R,et al.2010.Enhanced drought-tolerance in the homoploid hybrid species Pinus densata:implication for its habitat divergence from two progenitors.New Phytologist,185(1):204-216.
Mallet J.2005.Hybridization as an invasion of the genome.Trends in Ecology&Evolution,20(5):229-237.
Mantgem P J V,Stephenson N L,Byrne J C,et al.2009.Widespread increase of tree mortality rates in the western United States.Science,323(5913):521-524.
Mao J F,Wang X R.2011.Distinct niche divergence characterizes the homoploid hybrid speciation of Pinus densata on the Tibetan plateau.American Naturalist,177(4):424-439.
Meinshausen M,Smith S J,Calvin K,et al.2011.The RCP greenhouse gas concentrations and their extensions from 1765 to 2300.Climatic Change,109(1):213-241.
Pearson R G,Dawson T P.2003.Predicting the impacts of climate change on the distribution of species:are bioclimate envelope models useful?Global Ecology&Biogeography,12(5):361-371.
Phillips S J,Anderson R P,Schapire R E.2008.Modeling of species distributions with Max Ent:new extensions and a comprehensive evaluation.Ecography,31(2):161-175.
R Development Core Team.2014.R:A language and environment for statistical computing.R foundation for statistical computing.Vienna,Austria.
Ren Z P,Wang D Q,Ma A M,et al.2016.Predicting malaria vector distribution under climate change scenarios in China:challenges for malaria elimination.Scientific Reports,6:20604.
Rieseberg L H,Archer M A,Wayne R K.1999.Transgressive segregation,adaptation and speciation.Heredity,83(4):363-372.
Rieseberg L H,Lexer C.2003.Major ecological transitions in wild sunflowers facilitated by hybridization.Science,301(5637):1211-1216.
Rieseberg L H.1997.Hybrid origins of plant species.Annual Review of Ecology and Systematics,28(1):359-389.
Rundle H D,Nosil P.2005.Ecological speciation.Ecology Letters,8(3):336-352.
Schumer M,Rosenthal G G,Andolfatto P.2014.How common is homoploid hybrid speciation?Evolution,68(6):1553-1560.
Schwarzbach A E,Donovan L A,Rieseberg L H.2001.Transgressive character expression in a hybrid sunflower species.American Journal of Botany,88(2):270-277.
Sun Y,Abbott R J,Li L,et al.2014.Evolutionary history of Purple cone spruce(Picea purpurea)in the Qinghai-Tibet Plateau:homoploid hybrid origin and Pleistocene expansion.Molecular Ecology,23(2):343-359.
Szewczyk J.2009.Tree mortality and effects of release from competition in an old-growth Fagus-Abies-Picea stand.Journal of Vegetation Science,12(5):621-626.
Wang B,Mao J,Gao J,et al.2011.Colonization of the Tibetan Plateau by the homoploid hybrid pine Pinus densata.Molecular Ecology,20(18):3796-3811.
Xu D,Yan H.2001.A study of the impacts of climate change on the geographic distribution of Pinus koraiensis in China.Environment International,27(2):201-205.
Xu Y,Xu C H.2012.Preliminary assessment of simulations of climate changes over China by CMIP5 multi-models.Atmospheric and Oceanic Science Letters,5(6):489-494.
Yakimowski S B,Rieseberg L H.2014.The role of homoploid hybridization in evolution:a century of studies synthesizing genetics and ecology.American Journal of Botany,101(8):1247-1258.