浙东晚中新世植物解剖学与有机地球化学综合分析
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摘要
新生代是被子植物迅速演化的一个重要时期，也是环境变化和现代气候系统形成、定型的关键时期。浙江东部上中新统下南山组含有丰富的压型植物化石，为分析化石植物的解剖学以及有机地球化学特征提供了条件。目前，古植物解剖学与有机地球化学不但是研究化石植物埋藏学特征，尤其是分子埋藏学的最先进的两个方法，同时又是古环境定量研究中应用最广泛的两个方法。
从下南山组植物群中选取四种优势属种作为本文研究的主要对象：Ilex cornuta、Quercus delavayi、Cunninghamia lanceolata和Liquidambar miosinica。运用埋藏学、植物解剖学、植物生理学和有机地球化学等学科交叉研究的方法对这四种化石进行了分析，获得了浙东化石植物的分子埋藏学数据和古环境信息。
开展下南山组四种植物的热解-气相色谱／质谱(Py-GC／MS)分析，指示了四种化石植物的有机分子保存情况(从好到差)：L.miosinic≧C.lanceolata＞I.cornuta＞Q.delavayi。Py-GC／MS总离子色谱图(TIC)展示出四种化石及其现生对应植物的脂肪族化合物的变化，并将它们与下南山组沉积物的脂肪族同系物进行比较，表明下南山组四种植物化石脂肪族分子的保存机制属于“内源性的原位聚合”(in situ polymerization)。
利用栎属植物的气孔指数对CO_2变化的敏感性，根据化石Q．delavayi的气孔指数值，分别采用气孔比率法和气孔指数法获得的浙东晚中新世的CO_2浓度非常接近，分别为321ppmv和338ppmv，该结果与同时代采用其他方法或其他样品获得的结果也接近。
从下南山被子植物群中鉴定出85个叶片形态型，采用叶缘分析法(LMA)获得了浙江东部天台晚中新世的年均温为19.7～22.9℃，将该值代入本文建立的年均温和年较差的回归关系方程，计算出天台晚中新世的年较差为12.9～17.5℃，由这两个参数推测下南山植物群属于亚热带向热带过渡的植被类型。
现生和化石枫香的角质层上都含有表皮毛／毛基，利用不同地区间现生枫香叶片上表皮毛的变化建立毛基密度(个／cm~2)与气候参数的关系，根据化石枫香的毛基密度(约10个／cm~2)指示了天台晚中新世的气候与现在广州相似。
根据经验公式将化石枫香的稳定碳同位素值换算成晚中新世大气中的稳定碳同位素值δ~(13)C_(atm)，将之与大洋钻探碳酸盐岩中获得的δ~(13)C_(atm)比较，表明化石枫香不适合作为推算晚中新世δ~(13)C_(atm)的材料。采集于美国Clarkia中新统地层中的δ~(13)C也同样指示了化石枫香不能作为推算晚中新世δ~(13)C_(atm)的可靠材料。校正后的化石枫香碳同位素分馏值(△~(13)C)受环境因素的综合影响，可能来自于光照、温度等，但是与大气CO_2浓度和水分没有关系或关系很小。利用枫香的△~(13)C恢复出中新世下南山的化石L.miosinica水分利用效率(WUE)约为82μmolCO_2／molH_2O，Clarkia的化石L.pachyphyllum约为97μmolCO_2／molH_2O，均比其现生种高，且阳叶的WUE均比阴叶高，表明光照、温度是枫香WUE变化的主要影响因素。
The Cenozoic is a very important era in which climatic and environmental dynamicsresemble modern ones, and angiosperms undergo intensely adaptive radiation. A Mioceneflora from the Xiananshan Formation of eastern Zhejiang Province, containing veryabundant compressive plant fossils, is a significant key for plant anatomy and organicgeochemical research. Now, anatomy and organic geochemistry are extremely usefulmethods and techniques for fossil taphonomy, molecular taphonomy and quantitativelypalaeoenvironmental reconstruction.
Four predominant species in the Xiananshan Formation, including llex cornuta, Quercusdelavayi, Cunninghamia lanceolata and Liquidambar miosinica, are chosen from theMiocene Xiananshan Flora as materials of the current study. The molecular taphonomicdata and palaeoenvironmental information of the fossil flora are derived from theinvestigations on the four species by application of related taphonomic, anatomic,physiologic and organic geochemic techniques.
Py-GC/MS results provide clues on organic molecular preservation status of the fourfossil species (from better to worse): L. miosinica≧C. lanceolata＞I.cornuta＞Q.delavayi. Variations of aliphatic compound observed on TIC among the four fossil species,and corresponding nearest living relative species (NLRs) as well as the deposits ofXiananshan Formation, show that aliphatic molecular preservation of the fossil remains isassigned to "in situ polymerization".
As is well-known to all, a close negative correlation exists between atmospheric CO_2concentration and stomatal index of Quercus. Therefore, based on stomatal ratios andstomatal indexes of fossil and extant leaves of Q. delavayi, palaeoatmospheric CO_2 levels inthe late Miocene of Tiantai are estimated as 321 ppmv and 338 ppmv, respectively. Theresults are consistent with other relevant studies.
85 morphotypes are identified from about 1000 angiosperm specimens of the XiananshanFlora. Consequently, the mean annual temperature (MAT) of 19.7～22.9℃in the LateMiocene Tiantai is derived from leaf margin analysis, the difference of temperatures ofcoldest and warmest mongth (DT) of 12.9～17.5℃is originated from a regressive correlation formula which is established between current MAT and DT. It indicates that theXiananshan flora belongs to a transitional type from subtropic to tropic vegetation.
A close correlation is established between trichome base density of modern L. formosanaand climatic parameters. The trichome base density of fossil L. miosinica is ca. 10 (cm~(-2)),indicating that the late Miocene climate in Tiantai is similar to that in nowaday Guangzhou.
Based on the empirical formula of Arens et al. (2000), atmospheric carbon isotopiccomposition (δ~(13)C_(atm))in the late Miocene is calculated from carbon isotopic composition(δ~(13)C) of L. miosinica and is compared toδ~(13)C_(atm) from carbonate isotope record, suggestingthat the species is not suitable forδ~(13)C_(atm) estimates. And the result based on the fossil L.pachyphyllum from the Miocene Clarkia Formation in the U.S.A. agrees with the viewpoint.Carbon isotopic discrimination (Δ~(13)C) of fossil Liquidambar is mainly related toenvironmental factors, such as irradiation, temperature, other than atmospheric CO_2concentration and water availability. According toΔ~(13)C of fossil leaves, water useefficiency (WUE) is estimated to be 82μmolCO_2/molH_2O for L. miosinica,97μmolCO_2/molH_2O for L. pachyphyllum, respectively, and both the values are higher thantheir NLRs as well as WUE of sun morphotypes higher, which indicates that sunlight ortemperature is the significant impact on the WUE.
从下南山组植物群中选取四种优势属种作为本文研究的主要对象：Ilex cornuta、Quercus delavayi、Cunninghamia lanceolata和Liquidambar miosinica。运用埋藏学、植物解剖学、植物生理学和有机地球化学等学科交叉研究的方法对这四种化石进行了分析，获得了浙东化石植物的分子埋藏学数据和古环境信息。
开展下南山组四种植物的热解-气相色谱／质谱(Py-GC／MS)分析，指示了四种化石植物的有机分子保存情况(从好到差)：L.miosinic≧C.lanceolata＞I.cornuta＞Q.delavayi。Py-GC／MS总离子色谱图(TIC)展示出四种化石及其现生对应植物的脂肪族化合物的变化，并将它们与下南山组沉积物的脂肪族同系物进行比较，表明下南山组四种植物化石脂肪族分子的保存机制属于“内源性的原位聚合”(in situ polymerization)。
利用栎属植物的气孔指数对CO_2变化的敏感性，根据化石Q．delavayi的气孔指数值，分别采用气孔比率法和气孔指数法获得的浙东晚中新世的CO_2浓度非常接近，分别为321ppmv和338ppmv，该结果与同时代采用其他方法或其他样品获得的结果也接近。
从下南山被子植物群中鉴定出85个叶片形态型，采用叶缘分析法(LMA)获得了浙江东部天台晚中新世的年均温为19.7～22.9℃，将该值代入本文建立的年均温和年较差的回归关系方程，计算出天台晚中新世的年较差为12.9～17.5℃，由这两个参数推测下南山植物群属于亚热带向热带过渡的植被类型。
现生和化石枫香的角质层上都含有表皮毛／毛基，利用不同地区间现生枫香叶片上表皮毛的变化建立毛基密度(个／cm~2)与气候参数的关系，根据化石枫香的毛基密度(约10个／cm~2)指示了天台晚中新世的气候与现在广州相似。
根据经验公式将化石枫香的稳定碳同位素值换算成晚中新世大气中的稳定碳同位素值δ~(13)C_(atm)，将之与大洋钻探碳酸盐岩中获得的δ~(13)C_(atm)比较，表明化石枫香不适合作为推算晚中新世δ~(13)C_(atm)的材料。采集于美国Clarkia中新统地层中的δ~(13)C也同样指示了化石枫香不能作为推算晚中新世δ~(13)C_(atm)的可靠材料。校正后的化石枫香碳同位素分馏值(△~(13)C)受环境因素的综合影响，可能来自于光照、温度等，但是与大气CO_2浓度和水分没有关系或关系很小。利用枫香的△~(13)C恢复出中新世下南山的化石L.miosinica水分利用效率(WUE)约为82μmolCO_2／molH_2O，Clarkia的化石L.pachyphyllum约为97μmolCO_2／molH_2O，均比其现生种高，且阳叶的WUE均比阴叶高，表明光照、温度是枫香WUE变化的主要影响因素。
The Cenozoic is a very important era in which climatic and environmental dynamicsresemble modern ones, and angiosperms undergo intensely adaptive radiation. A Mioceneflora from the Xiananshan Formation of eastern Zhejiang Province, containing veryabundant compressive plant fossils, is a significant key for plant anatomy and organicgeochemical research. Now, anatomy and organic geochemistry are extremely usefulmethods and techniques for fossil taphonomy, molecular taphonomy and quantitativelypalaeoenvironmental reconstruction.
Four predominant species in the Xiananshan Formation, including llex cornuta, Quercusdelavayi, Cunninghamia lanceolata and Liquidambar miosinica, are chosen from theMiocene Xiananshan Flora as materials of the current study. The molecular taphonomicdata and palaeoenvironmental information of the fossil flora are derived from theinvestigations on the four species by application of related taphonomic, anatomic,physiologic and organic geochemic techniques.
Py-GC/MS results provide clues on organic molecular preservation status of the fourfossil species (from better to worse): L. miosinica≧C. lanceolata＞I.cornuta＞Q.delavayi. Variations of aliphatic compound observed on TIC among the four fossil species,and corresponding nearest living relative species (NLRs) as well as the deposits ofXiananshan Formation, show that aliphatic molecular preservation of the fossil remains isassigned to "in situ polymerization".
As is well-known to all, a close negative correlation exists between atmospheric CO_2concentration and stomatal index of Quercus. Therefore, based on stomatal ratios andstomatal indexes of fossil and extant leaves of Q. delavayi, palaeoatmospheric CO_2 levels inthe late Miocene of Tiantai are estimated as 321 ppmv and 338 ppmv, respectively. Theresults are consistent with other relevant studies.
85 morphotypes are identified from about 1000 angiosperm specimens of the XiananshanFlora. Consequently, the mean annual temperature (MAT) of 19.7～22.9℃in the LateMiocene Tiantai is derived from leaf margin analysis, the difference of temperatures ofcoldest and warmest mongth (DT) of 12.9～17.5℃is originated from a regressive correlation formula which is established between current MAT and DT. It indicates that theXiananshan flora belongs to a transitional type from subtropic to tropic vegetation.
A close correlation is established between trichome base density of modern L. formosanaand climatic parameters. The trichome base density of fossil L. miosinica is ca. 10 (cm~(-2)),indicating that the late Miocene climate in Tiantai is similar to that in nowaday Guangzhou.
Based on the empirical formula of Arens et al. (2000), atmospheric carbon isotopiccomposition (δ~(13)C_(atm))in the late Miocene is calculated from carbon isotopic composition(δ~(13)C) of L. miosinica and is compared toδ~(13)C_(atm) from carbonate isotope record, suggestingthat the species is not suitable forδ~(13)C_(atm) estimates. And the result based on the fossil L.pachyphyllum from the Miocene Clarkia Formation in the U.S.A. agrees with the viewpoint.Carbon isotopic discrimination (Δ~(13)C) of fossil Liquidambar is mainly related toenvironmental factors, such as irradiation, temperature, other than atmospheric CO_2concentration and water availability. According toΔ~(13)C of fossil leaves, water useefficiency (WUE) is estimated to be 82μmolCO_2/molH_2O for L. miosinica,97μmolCO_2/molH_2O for L. pachyphyllum, respectively, and both the values are higher thantheir NLRs as well as WUE of sun morphotypes higher, which indicates that sunlight ortemperature is the significant impact on the WUE.
引文
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