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气候变化对植物分布与多样性的影响 Climate Change Impacts on Plant Distribution and Diversity Zhiheng Wang (王志恒) [email protected] 15.05.2014 @ PKU, Beijing Department of Ecology College of Urban and Environmental Sciences Peking University Outline 1. 2. 3. 4. What’s macroecology (宏观生态学) Deep-time climate changes and plant evolution Impacts of climate changes since the LGM on plant diversity Threats of global warming in 20th and 21st centuries on plant diversity 一、宏观生态学(macroecology) Science 1989 Central question: mechanisms of patterns in life distribution and diversity in space and time Scale: regional –> global Methods: hypothesis testing + quantitative statistics + comparative phylogenetics 1995 James H. Brown Macroecology is young, but it is one of most active sub-disciplines of ecology. 宏观生态学是生态学中年轻、但最活跃的分支之一 宏观生态学(macroecology) Journal of Biogeography (IF: 4.9) Ecography (IF: 5.1) Global Ecology & Biogeography (IF: 7.2) Diversity & Distributions (IF: 6.1) 影响因子 Major Journals for macroecology: 年份 Beck et al. 2012 宏观生态学 生物地理学 学科基础 生态学 地理学 研究内容 物种分布和多样性格局的形成机制 物种和多样性的地理格局 生物分布与环境和进化的关系 生物分布的格局与样式 定量 定性 侧重点 方法 Global climate changes Sea surface temperature changes Eocene climatic optimum (55 ma) Mid-Cretaceous Greenhouse (90-100 ma) Clarke et al. 1999. Geology Global climate changes Quaternary (2.3 ma) Age (Ma) Eocene-Oligocene transition (34 ma) Eocene climatic optimum (55 ma) Temperature (℃) Zachos et al. 2001. Science, 292, 686-693 Climate change impact on species diversity Long-term geological history 1-100 Ma Long-term geological history < 1 Ma Speciation Extinction Dispersal √ √ √ √ √ industrial 1-100 √ evolution years • Fossil records and paleontological methods Short-term current 1-100 √ century years • DNA data and comparative phylogenetic methods Short-term √ √ Deep-time climatic optimum and plant diversification Global speciation rate of plants based on fossil evidence Crepet et al. 2009. Am J Bot extinct sp. new sp. Deep-time climatic optimum and plant diversification Jaramillo et al. 2010. Science Eocene-Oligocene transition Age (Ma) Eocene-Oligocene transition (34 ma) Temperature (℃) Zachos et al. 2001. Science, 292, 686-693 Eocene-Oligocene transition (ca. 34 Ma) Eocene-Oligocene transition (ca. 34 Ma) Zanazzi et al. 2007 Nature Effects of Eocene-Oligocene transition Global mammal extinction Fossil records of North European mammals 33.4 Ma young old Eocene–Oligocene extinction event Grande Coupure (great break) Hooker et al. 2004. J Geolog Soc Effects of Eocene-Oligocene transition Plant molecular evolutionary rate Rhododendron Quercus Evolution of Rhododendron Zhiheng Wang, Xiaoting Xu, Peking Univ., China Dimitar Dimitra, Oslo Univ., Norway Alexandre Antonelli, Univ. of Gothenburg, Sweden Katsuhiro Nakao, Forestry and Forest Products Research Institute, Japan Alexandra Muellner-Riehl, Univ. of Leipzig, Germany Biogeography of Rhododendron Species diversity: c.a. 900 sp; c.a. 550 sp in China Distribution: Northern Hemisphere Diversification Originated in late Cretaceous to early Paleogene (50-70 mya) Biogeography of Rhododendron Sequence data from GenBank 388 species 16 genes: atpB-rbcL, rbcL, matK, ndhF, psbA-trnH, trnL-F, trnL, trnT-trnL, trnStrnG, ITS, RPB2I-1, RPB2I-2, RPB2I-3, RPB2I-4, RPB2I-5, RPB2I-6 Substitution rate Evolutionary rate of Rhododendorn Age (Ma) Biogeography of Rhododendron Question: What’s the mechanism of the high Rhododendron species diversity? Hypothesis: 1) It has been believed that the rapid diversification of Rhododendron was enhanced by the rise of Tibetan Plateau at ca. 30 – 40 Ma. Age (Ma) 2) Global climate shifted from Eocene greenhouse to Oligocene icehouse at 34 Ma. Cool climate led to the expansion of temperate vegetation, and then the diversification of Rhododendron. EoceneOligocene transition (34 ma) Temperature (℃) Biogeography of Rhododendron Expectation of Hypothesis 1 Evolutionary rate is high during the period of the collision between Indian subcontinent and Eurasia, and the clades originated in association with the collision (those in southwest China) have high evolutionary rate. Expectation of Hypothesis 2 Evolutionary rate is high during the period of the collision between Indian subcontinent and Eurasia, and the clades experienced high climatic changes (those in the north) have high evolutionary rate. Biogeography of Rhododendron Substitution rate Molecular evolutionary rate of different clades Clade 1 Clade 2 Clade 3 Clade 4 34 Ma Age (Ma) Clade 5 Clade 6 Clade 7 Evolution of Quercus Xiaoting Xu, Peking Univ., China Dimitar Dimitra, Oslo Univ., Norway Global pattern of oak (Quercus) diversity 北温带森林的优势树种 ca. 450 spp., 407 spp. included 186本植物志、数据库和发表文献 进化速率与栎属物种多样性分化 分子进化速率的时间变化 序列 序列alignment:mafft -一致性序列构建(consensus sequences) 进化树构建 Maximum likelihood method RAxML-HPC version 7.4.2 定年 Beast 1.7.4 化石校正 Supermatrix: 11 genes, 40 × 9528 Gaps:小于50% linked Clock model 栎属分子进化速率的时间变化 11 genes 40 sp.× 9528 bp 缺失数据:<50% BESAT: Link Clock model Climate change impact on species diversity Speciation Extinction Dispersal √ √ √ Long-term geological history 1-100 Ma Long-term geological history < 1 Ma √ √ Short-term industrial evolution 1-100 years √ √ Short-term current climate 1-100 √ Quaternary change century yearsthe Last Glacial Maximum Climate change since √ Modeled distribution and population size of megafauna species at 42, 30, 21 and 6 kyr BP. Effective population size Effective population size was estimated by population genetic methods based on ancient DNA. Lorenzen, et al. 2011. Nature Influences of climate change since the LGM on Chinese woody plant diversity Yaoqi Li Xiaoting Xu Zhiheng Wang Data of plant distributions From “Database of China’s Woody Plants (v2.0)” ● compiled from more than 320 national and provincial floras, many local floras and specimen records ● examined by 21 local experts of plants ● c.a. 6 years ● Taxonomy: Flora of China (English version) ● Specimen records + observation data ● Species number: 11405 Specimen records observation data Climate change impacts on Chinese plant diversity Anomaly of mean annual temperature since the LGM Modern climate Mean winter temperature Anomaly = LGM MAT – Modern MAT Precipitation Temperature LGM climate data was hindcasted by Community Climate System Model (CCSM). Climate change impacts on Chinese plant diversity Method: Geographically weighted regression (GWR) Local R2 of temperature anomaly Local R2 of modern climate Climate change impacts on Chinese plant diversity R2 difference Anomaly – MAT Anomaly – winter T Anomaly – MAP Temperature change since the LGM explains more richness variation than modern climate in southeastern China. Principal component analysis of modern climate Energy: MAT + Bio4(Temperature Seasonality ) + PET R2 of water PC 1 R2 of energy PC 1 Water: MAP + Bio15(Precipitation Seasonality) + AET Energy PC 1 Anomaly 80.22% energy PCof1total variance Water PC 1 86.67% of Anomaly - total water PC 1 variance Temperature change since the LGM explains more richness variation than modern climate in southeastern China. Influences of climate change since the LGM on Chinese vegetation Siyang Wang Zhiheng Wang Data: 1) 1:1,000,000 vegetation map 2) LGM climate data was hindcasted by Community Climate System Model (CCSM). Method: species distribution models LGM & pollen data steppe Broadleaved evergreen/warm mixed forest desert temperate deciduous forest taiga Modern LGM Transformation LGM V1 V2 V3 V4 V5 V6 V7 V8 V9 V10 V11 V12 V13 V14 V15 V16 V17 V18 V19 V20 V21 V22 modern 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 >=40% >=10% <40% <10% * Substantial contraction: tropical rainforest and monsoon forest, temperate needleleaf-broadleaf mixed forest, temperate deciduous scrub Forests move northward since the LGM modern distrib LGM distrib. LGM distrib. Veget1: Cool and temperate coniferous forest Veget6: Temperate deciduous forest Steppe and desert contracted Veget16: temperate steppe veget22: alpine tundra Climate change impact on species diversity Speciation Dispersal √ √ < 1 Ma √ √ industrial evolution 1-100 years √ √ current century 1-100 years √ √ Long-term geological history 1-100 Ma Long-term geological history Short-term Short-term √ Extinction Recent climate change IPCC report V, 2013 Recent climate change IPCC report V, 2013 Plants track climate change period 1: 1905-1985 period 2: 1985-2005 Lenoir et al. 2008. Science Plant species move upward along French Alpine Speed: 29 m/decade Plants track climate change Plant traits affect moving speed Lenoir et al. 2008. Science Plants track climate change Spider Butterfly Ground beetles Grasshopper Northern boundary of species distribution moves northward in UK. Speed: 16.9 km/decade Chen et al. 2011. Science Species diversity changes in Changbai Mts. Time period: 1963 – 2006 Species diversity declined at the same environment. Climate change impact on species diversity Speciation Dispersal √ √ < 1 Ma √ √ industrial evolution 1-100 years √ √ current century 1-100 years √ √ Long-term geological history 1-100 Ma Long-term geological history Short-term Short-term √ Extinction 5 Ongoing project Threats of climate change on woody plant diversity Global temperature change Source: National Climatic Data Center, US China temperature change Source: Wang et al. 2010 Global warming changes the habitat, growth, phenology and distribution ranges of organisms, and may lead to migration or extinction How climate change influences plants in China ? Current and future climate data Source and variables • • • From the WorldClim website Resolution: 2.5 × 2.5 arc min Variables: Bio1 – Bio19 Current climate Precipitation Temperature Scenarios of future climate • • • A1B: maximum energy requirements, balance across fuel sources A2: high energy requirements B2: lower energy requirements Climate in 2080 (A2) Climate change Model calibration Models 1) Generalized linear model (GLM) with binomial residuals 2) Maximum entropy (Maxent) 3) Classification tree (CT) Variable selection 1) Stepwise regression using GLM for a species Si: forward + backward 2) Select a subset of variables for Si based on Akaike information criterion (AIC) 3) The selected subset of variables were used for Maxent and CT Species selection Species with range size > 20 grid cells 7437 species in total modeled species all species Vulnerable vs. benefited areas Present richness patterns (1950 – 2000) Projections in 2080 – 2100 A1B A2 最大能量消耗 能源均衡发展 高能量消耗 B2 低能量消耗 Projected species richness patterns Results 1. Vulnerable areas: central to east and south China 2. Benefited areas: Tibetan Plateau Wang el al. in preparation Changes in species richness Decline No change Increase Projected species movement Elevation change Latitude change Predicted species dispersal North Daxing’an Present North Taihang Future Changbai Kunlun-Qilian Wuyi Hengduan Based on the second threshold method Acknowledgements Prof. Jingyun Fang Prof. Bernhard Schmid Prof. Carsten Rahbek Dr. Xiaoting Xu Dr. Zhiyao Tang Dr. Yining Liu Dr. Xiujuan Qiao Dr. Zhaodi Guo Dr. Luying Tang The 21 experts who reviewed the species distribution data Thank you for your attention Two periods of climatic optimum Eocene climatic optimum (55 ma) Mid-Cretaceous Greenhouse (90-100 ma) Bowen et al. 2004. Science