地质时期樟科植物花化石及其系统演化意义
|
摘要
该研究对地质时期樟科植物花化石的主要类群,即Androglandula、Lauranthus、Mauldinia、Neusenia、Perseanthus和Potomacanthus属分别从属和种的形态特征、分布、地层以及系统意义进行了论述,并结合现代樟科植物从地层和分布、花序类型、花部形态特征和显微构造特征进行了分析。结果表明:(1)樟科植物在中晚白垩纪期,已经起源于劳亚古陆的中纬度区域。(2)樟科植物的花序类型为Mauldinia属的侧生花序类型和假伞形花序类型。(3)花为3基数的两性花,花被片6枚排列为2轮,雄蕊12或6,排列为4、3或2轮,最内轮雄蕊不育,第三轮雄蕊基部常见一对附属腺体,雄蕊药室瓣裂,4或2药室,雌蕊为单心皮。(4)花被片上常有大量的油细胞、并列型气孔器和单细胞毛。该研究结果中樟科花化石的发现,为樟科植物的系统演化提供了古生物学的证据和资料。
This article systematically reviews the fossil record of the Lauraceae flower,including Androglandula,Lauranthus,Mauldinia,Neusenia,Perseanthus and Potomacanthus which were found in the geological times,the morphological characters,distribution,stratum and phylogenetic significance of these genera and species are systematically discussed. This article explores the stratum,distribution,inflorescence types,floral morphological characters,microstructural characters with the modern Lauraceae. The results were as follows:( 1) Lauraceae originated in the mid-latitude region of the Laurasia from middle to late Cretaceous.( 2) Two inflorescences are found in the fossil flower of Lauraceae,one is Mauldinia type,which has elongated axes bearing distinctive,spirally arranged lateral,bilobed cladode-like units with five sessile bisexual flowers on the adaxial surface,the other is pseudoumbel type which has three sessile and closely spaced flowers with enclosed by two broad and strongly truncate bracets.( 3) Bisexual flowers with three cardinal numbers,six perianth lobs arranging in two whorls,twelve or six stamens in four,three or two whorls,innermost whorl reduced to staminodes,the third whorl stamen usually bearing two sessile and distinct basal glandular protuberances,anthers have two or four valved chamber,the gynoecium has single carpel.( 4) A large number of oil cells,paracytic stomatal apparatus and unicellular hairs are often present on perianth. The flower fossils of Lauraceae provide geological and historical evidences for the phylogenetic evolutionary study of the Lauraceae.
This article systematically reviews the fossil record of the Lauraceae flower,including Androglandula,Lauranthus,Mauldinia,Neusenia,Perseanthus and Potomacanthus which were found in the geological times,the morphological characters,distribution,stratum and phylogenetic significance of these genera and species are systematically discussed. This article explores the stratum,distribution,inflorescence types,floral morphological characters,microstructural characters with the modern Lauraceae. The results were as follows:( 1) Lauraceae originated in the mid-latitude region of the Laurasia from middle to late Cretaceous.( 2) Two inflorescences are found in the fossil flower of Lauraceae,one is Mauldinia type,which has elongated axes bearing distinctive,spirally arranged lateral,bilobed cladode-like units with five sessile bisexual flowers on the adaxial surface,the other is pseudoumbel type which has three sessile and closely spaced flowers with enclosed by two broad and strongly truncate bracets.( 3) Bisexual flowers with three cardinal numbers,six perianth lobs arranging in two whorls,twelve or six stamens in four,three or two whorls,innermost whorl reduced to staminodes,the third whorl stamen usually bearing two sessile and distinct basal glandular protuberances,anthers have two or four valved chamber,the gynoecium has single carpel.( 4) A large number of oil cells,paracytic stomatal apparatus and unicellular hairs are often present on perianth. The flower fossils of Lauraceae provide geological and historical evidences for the phylogenetic evolutionary study of the Lauraceae.
引文
BANDULSKA H,1926.On the cuticles of some fossil and recent Lauraceae[J].Bot J Linn Soc,47:383-424.
BANDULSKA H,1929.Secretory cells in a fossil leaf[J].Ann Bot,43:203-204.
BALTHAZAR MV,PEDERSEN KP,CRANE PR,et al,2007.Potomacanthus lobatus gen.et sp.nov.,a new flower of probable Lauraceae from the Early Cretaceous(Early to Middle Albian)of Eastern North America[J].Am J Bot,94(12):2041-2053.
CHU QG,HU ZH,1999.Comparative anatomy of oil cells and mucilage cells in the leaves of the Lauraceae in China[J].Acta Phytotax Sin,37(6):529-540.[初庆刚,胡正海,1999.中国樟科植物叶中油细胞和粘液细胞的比较解剖研究[J].植物分类学报,37(6):529-540.]
CUI HB,1987.A study on the system of Lindera[J].Acta Phytotax Sin,25(3):161-171.[崔鸿宾,1987.山胡椒属系统的研究[J].植物分类学报,25(3):161-171.]
DAO KQ,XIE SP,SUN BN,et al,2011.Leaf morphology and cuticular microstructure of Cinnamomum cf.camphora from the Pliocene of Tengchong,Yunnan[J].Acta Palaeontol Sin,50(2):240-250.[刀克群,解三平,孙柏年,等,2011.云南腾冲上新统Cinnamomum cf.camphora叶形态和表皮微细构造研究[J].古生物学报,50(2):240-250.]
DICHER,1974.Approaches to the identification of angiosperm leaf remains[J].Bot Rev,40(1):1-157.
DRINNAN AN,CRANE PR,FRIIS EM,et al,1990.Lauraceous flowers from the Potomac Group(mid-Cretaceous)of eastern North America[J].Bat Gaz,151(3):370-384.
EKLUND H,2000.Lauraceous flowers from the late Cretaceous of North Carolina,USA[J].Bot J Linn Soc,132:397-428.
EKLUND H,KAVACEK J,1998.Lauraceous inflorescences and flowers from the Cenomanian of Bohemia(Czech Republic,central Europe)[J].Int J Plant Sci,159(4):668-686.
FRUMIN S,EKLUND H,FRIIS EM,2004.Mauldinia hirsuta sp.nov.,a new member of the extinct genus Mauldinia(Lauraceae)from the late Cretaceous(Cenomanian-Turonian)of Kazakhstan[J].Int J Plant Sci,165(5):883-895.
HERENDEEN PS,CREPET WL,NIXON KC,1994.Fossil flowers and pollen of Lauraceae from the Upper Cretaceous of New Jersey[J].Plant Syst Evol,189:29-40.
HERENDEEN PS,MAGALLON-PUEBLA S,LUPIA R,et al,1999.A preliminary conspectus of the Allon flora from the Late Cretaceous(late Santonian)of central Georgia,USA[J].Ann Mo Bot Gard,86(2):407-471.
HILL R,1986.Lauraceous leaves from the Eocene of Nerriga,New South Wales[J].Alcheringa,10(4):327-351.
HUANG JF,LI L,WERFF H,et al,2016.Origins and evolution of cinnamon and camphor:A phylogenetic and historical biogeographical analysis of the Cinnamomum group(Lauraceae)[J].Mol Phylogenet Evol,96:33-44.
LI J,LI XW,2004.Advanses in Lauraceae systematic research on the world scale[J].Acta Bot Yunnan,26(1):1-11[李捷,李锡文,2004.世界樟科植物系统学研究进展[J].云南植物研究,26(1):1-11.]
LI J,CHRISTOPHEL DC,CONRAN JG,et al,2004.Phylogenetic relationships within the‘core’Laureae(Litsea complex,Lauraceae)inferred from sequences of the chloroplast gene mat K and nuclear ribosomal DNA ITS regions[J].Plant Syst Evol,26(1):19-36.
LI JZ,QIU J,LIAO WB,et al,2009.Eocene fossil Alseodaphne from Hainan Island of China and its paleoclimatic implications[J].Sci Chin Ser D,52(10):1537-1542.
LI JZ,2010.Eocene Lauraceae plants from Changchang Basin,Hainan Island of China and its paleoenvironment significance[D].Guangzhou:Sun Yat-Sen University:1-103.[李景照,2010.海南岛长昌盆地始新世樟科植物及古环境意义[D].广州:中山大学:1-103.]
LI XW,BAI PY,LI YR,et al,1982.Flora Republicae Popularis Sinicae[M].Beijing:Science Press,31:1-465.[李锡文,白佩瑜,李雅茹,等,1982.中国植物志[M].北京:科学出版社,31:1-465.]
LI XW,1995.The origin and evolution of Litsea Genera Group(Laureae)in Lauraceae[J].Acta Bot Yunnan,17(3):251-254.[李锡文,1995.樟科木姜子属群的起源与演化[J].云南植物研究,17(3):251-254.]
LI ZM,LI J,LI XW,2006.Phylogenetic of the genus Actinodaphne Nees(Lauraceae)inferred from the analyses of nr DNA ITS and ETS sequences[J].Acta Phytotax Sin,44(3):272-285.
LI ZM,2005.A phylogenetic study on the genus Actinodaphne Nees(Lauraceae)inferred from the morphological and molecular systematic data[D].Kunming:Xishuangbanna Tropical Botanical Garden,Chinese Academy of Sciences:1-74.[李志明,2005.基于形态学和分子系统学资料的樟科黄肉楠属系统学研究[D].昆明:中国科学院西双版纳热带植物园:1-74.]
LIU YS,1990.Cuticular studies on two Pleistocene species of Lauraceae in Baise Basin,Guangxi[J].Acta Bot Sin,32(10):805-808.[刘裕生,1990.广西百色盆地更新世樟科两种植物角质层研究[J].植物学报,32(10):805-808.]
MOREAU JD,GOMEZ B,DAVIERO-GOMEZ V,et al,2016.Inflorescences of Mauldinia sp.(Lauraceae)and associated fruits from the Cenomanian of Languedoc Roussillon,France[J].Cretac Res,59:18-29.
POLE M,2007.Lauraceae macrofossils and dispersed cuticle from the Miocene of Southern New Zealand[J].Palaeontol Electr,10(1)3A:38p.
RAVEN PH,AXELROD DI,1974.Angiosperm biogeographay and past continental movements[J].Ann Mo Bot Gard,61:539-673.
ROHWER JG,2000.Toward a phylogenetic classification of the Lauraceae:Evdence from mat K sequences[J].Syst Bot,25(1):60-71.
TAKAHASHI M,CRANE PR,ANDO H,1999.Fossil flower and associated plant fossils from the Kamikitaba locality(Ashizawa Formation,Futaba Group,Lower Coniacian,Upper Cretaceous)of Northeast Japan[J].J Plant Res,112:187-206.
TAKAHASHI M,HERENDEEN PS,CRANE PR,2001.Lauraceous fossil flower from the Kamikitaba locality(Lower Coniacian;Upper Cretaceous)in northeastern Japan[J].JPlant Res,114(4):429-434.
TAYLOR DW,1988.Eocene floral evidence of Lauraceae:corroboration of the North American megafossil record[J].Amer J Bot,75(7):948-957.
VIEHOFEN A,HARTKOPF-FRODER C,FRIIS EM,2008.Inflorescences and flowers of Mauldinia angustiloba sp.nov.(Lauraceae)from middle Cretaceous karst infillings in the Rhenish Massif,Germany[J].Int J Plant Sci,169(7):871-889.
WANG XJ,XIAO LT,DONG AW,et al,2017.Advances in plant science in China in 2016[J].Chin Bull Bot,52(4):394-452.[王小箐,萧浪涛,董爱武,等,2017.2016年中国植物科学若干领域重要研究进展[J].植物学报,52(4):394-452.]
WU ZY,LU AM,TANG YC,et al,2003.The families and genera of angiosperms in China,a comprehensive analysis[M].Beijing:Science Press:1-1209.[吴征镒,路安民,汤彦承,等,2003.中国被子植物科属综论[M].北京:科学出版社:1-1209.]
BANDULSKA H,1929.Secretory cells in a fossil leaf[J].Ann Bot,43:203-204.
BALTHAZAR MV,PEDERSEN KP,CRANE PR,et al,2007.Potomacanthus lobatus gen.et sp.nov.,a new flower of probable Lauraceae from the Early Cretaceous(Early to Middle Albian)of Eastern North America[J].Am J Bot,94(12):2041-2053.
CHU QG,HU ZH,1999.Comparative anatomy of oil cells and mucilage cells in the leaves of the Lauraceae in China[J].Acta Phytotax Sin,37(6):529-540.[初庆刚,胡正海,1999.中国樟科植物叶中油细胞和粘液细胞的比较解剖研究[J].植物分类学报,37(6):529-540.]
CUI HB,1987.A study on the system of Lindera[J].Acta Phytotax Sin,25(3):161-171.[崔鸿宾,1987.山胡椒属系统的研究[J].植物分类学报,25(3):161-171.]
DAO KQ,XIE SP,SUN BN,et al,2011.Leaf morphology and cuticular microstructure of Cinnamomum cf.camphora from the Pliocene of Tengchong,Yunnan[J].Acta Palaeontol Sin,50(2):240-250.[刀克群,解三平,孙柏年,等,2011.云南腾冲上新统Cinnamomum cf.camphora叶形态和表皮微细构造研究[J].古生物学报,50(2):240-250.]
DICHER,1974.Approaches to the identification of angiosperm leaf remains[J].Bot Rev,40(1):1-157.
DRINNAN AN,CRANE PR,FRIIS EM,et al,1990.Lauraceous flowers from the Potomac Group(mid-Cretaceous)of eastern North America[J].Bat Gaz,151(3):370-384.
EKLUND H,2000.Lauraceous flowers from the late Cretaceous of North Carolina,USA[J].Bot J Linn Soc,132:397-428.
EKLUND H,KAVACEK J,1998.Lauraceous inflorescences and flowers from the Cenomanian of Bohemia(Czech Republic,central Europe)[J].Int J Plant Sci,159(4):668-686.
FRUMIN S,EKLUND H,FRIIS EM,2004.Mauldinia hirsuta sp.nov.,a new member of the extinct genus Mauldinia(Lauraceae)from the late Cretaceous(Cenomanian-Turonian)of Kazakhstan[J].Int J Plant Sci,165(5):883-895.
HERENDEEN PS,CREPET WL,NIXON KC,1994.Fossil flowers and pollen of Lauraceae from the Upper Cretaceous of New Jersey[J].Plant Syst Evol,189:29-40.
HERENDEEN PS,MAGALLON-PUEBLA S,LUPIA R,et al,1999.A preliminary conspectus of the Allon flora from the Late Cretaceous(late Santonian)of central Georgia,USA[J].Ann Mo Bot Gard,86(2):407-471.
HILL R,1986.Lauraceous leaves from the Eocene of Nerriga,New South Wales[J].Alcheringa,10(4):327-351.
HUANG JF,LI L,WERFF H,et al,2016.Origins and evolution of cinnamon and camphor:A phylogenetic and historical biogeographical analysis of the Cinnamomum group(Lauraceae)[J].Mol Phylogenet Evol,96:33-44.
LI J,LI XW,2004.Advanses in Lauraceae systematic research on the world scale[J].Acta Bot Yunnan,26(1):1-11[李捷,李锡文,2004.世界樟科植物系统学研究进展[J].云南植物研究,26(1):1-11.]
LI J,CHRISTOPHEL DC,CONRAN JG,et al,2004.Phylogenetic relationships within the‘core’Laureae(Litsea complex,Lauraceae)inferred from sequences of the chloroplast gene mat K and nuclear ribosomal DNA ITS regions[J].Plant Syst Evol,26(1):19-36.
LI JZ,QIU J,LIAO WB,et al,2009.Eocene fossil Alseodaphne from Hainan Island of China and its paleoclimatic implications[J].Sci Chin Ser D,52(10):1537-1542.
LI JZ,2010.Eocene Lauraceae plants from Changchang Basin,Hainan Island of China and its paleoenvironment significance[D].Guangzhou:Sun Yat-Sen University:1-103.[李景照,2010.海南岛长昌盆地始新世樟科植物及古环境意义[D].广州:中山大学:1-103.]
LI XW,BAI PY,LI YR,et al,1982.Flora Republicae Popularis Sinicae[M].Beijing:Science Press,31:1-465.[李锡文,白佩瑜,李雅茹,等,1982.中国植物志[M].北京:科学出版社,31:1-465.]
LI XW,1995.The origin and evolution of Litsea Genera Group(Laureae)in Lauraceae[J].Acta Bot Yunnan,17(3):251-254.[李锡文,1995.樟科木姜子属群的起源与演化[J].云南植物研究,17(3):251-254.]
LI ZM,LI J,LI XW,2006.Phylogenetic of the genus Actinodaphne Nees(Lauraceae)inferred from the analyses of nr DNA ITS and ETS sequences[J].Acta Phytotax Sin,44(3):272-285.
LI ZM,2005.A phylogenetic study on the genus Actinodaphne Nees(Lauraceae)inferred from the morphological and molecular systematic data[D].Kunming:Xishuangbanna Tropical Botanical Garden,Chinese Academy of Sciences:1-74.[李志明,2005.基于形态学和分子系统学资料的樟科黄肉楠属系统学研究[D].昆明:中国科学院西双版纳热带植物园:1-74.]
LIU YS,1990.Cuticular studies on two Pleistocene species of Lauraceae in Baise Basin,Guangxi[J].Acta Bot Sin,32(10):805-808.[刘裕生,1990.广西百色盆地更新世樟科两种植物角质层研究[J].植物学报,32(10):805-808.]
MOREAU JD,GOMEZ B,DAVIERO-GOMEZ V,et al,2016.Inflorescences of Mauldinia sp.(Lauraceae)and associated fruits from the Cenomanian of Languedoc Roussillon,France[J].Cretac Res,59:18-29.
POLE M,2007.Lauraceae macrofossils and dispersed cuticle from the Miocene of Southern New Zealand[J].Palaeontol Electr,10(1)3A:38p.
RAVEN PH,AXELROD DI,1974.Angiosperm biogeographay and past continental movements[J].Ann Mo Bot Gard,61:539-673.
ROHWER JG,2000.Toward a phylogenetic classification of the Lauraceae:Evdence from mat K sequences[J].Syst Bot,25(1):60-71.
TAKAHASHI M,CRANE PR,ANDO H,1999.Fossil flower and associated plant fossils from the Kamikitaba locality(Ashizawa Formation,Futaba Group,Lower Coniacian,Upper Cretaceous)of Northeast Japan[J].J Plant Res,112:187-206.
TAKAHASHI M,HERENDEEN PS,CRANE PR,2001.Lauraceous fossil flower from the Kamikitaba locality(Lower Coniacian;Upper Cretaceous)in northeastern Japan[J].JPlant Res,114(4):429-434.
TAYLOR DW,1988.Eocene floral evidence of Lauraceae:corroboration of the North American megafossil record[J].Amer J Bot,75(7):948-957.
VIEHOFEN A,HARTKOPF-FRODER C,FRIIS EM,2008.Inflorescences and flowers of Mauldinia angustiloba sp.nov.(Lauraceae)from middle Cretaceous karst infillings in the Rhenish Massif,Germany[J].Int J Plant Sci,169(7):871-889.
WANG XJ,XIAO LT,DONG AW,et al,2017.Advances in plant science in China in 2016[J].Chin Bull Bot,52(4):394-452.[王小箐,萧浪涛,董爱武,等,2017.2016年中国植物科学若干领域重要研究进展[J].植物学报,52(4):394-452.]
WU ZY,LU AM,TANG YC,et al,2003.The families and genera of angiosperms in China,a comprehensive analysis[M].Beijing:Science Press:1-1209.[吴征镒,路安民,汤彦承,等,2003.中国被子植物科属综论[M].北京:科学出版社:1-1209.]