云南腾冲上新统Litsea cf.chunii的角质层微细构造特征
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摘要
在云南省腾冲县上新统芒棒组发现了保存较好的被子植物压型化石高山木姜子(相似种)Litseacf.chunii Cheng.该化石叶片呈窄椭圆形,基部楔形,顶端短尖,叶全缘,中脉直,二级脉羽状,与中脉夹角为30°~50°,近叶边缘微弯曲,羽状脉。进一步分析了化石叶片的角质层微细构造,利用叶结构结合表皮特征的分析方法提高了对化石植物分类鉴定的准确性。化石叶片的角质层特征主要表现为:上表皮无气孔器,表皮细胞多为四或五边形,排列呈规则的网格状,垂周壁直或微弯,平周壁光滑,毛基较少;下表皮细胞排列同上表皮,具平列型气孔器,其长轴无定向性,气孔器由下陷的保卫细胞和两个上拱的副卫细胞组成,具有单细胞毛基,根部由5~6个辐射状的细胞组成。同现生高山木姜子Litseachunii相比,化石叶片具有更小的毛基密度,推测毛基的增加是由于气候变冷所致。由此通过化石和现生叶片角质层的对比分析推断滇西地区上新世气候比现在温暖湿润。
A fossil leaf identified as Litsea cf.chunii Cheng was discovered in the late pliocene sediments of the Mangbang formation in Tengchong county,Yunnan province,China,which was well preserved with gross morphological structure and microstructure of cuticles in situ.The fossil leaf is narrowly elliptic,base cuneate, with obtuse apex and entire leaf margin.Its primary vein is straight and moderately thick;secondary veins are pinnate,which arise from the primary vein at angles from 30°~50°,and they are slightly curved near the leaf margin.Additionally,the anatomical characteristics of the present specimen are investigated as follows: the upper epidermis is free from stomata,composed of mostly tetragonal,pentagonal or slightly elongated cells,which are arranged in regular areola;anticlinal cell walls are straight and moderately thick,periclinal walls are smooth and slightly concave;trichome bases are rarely found.The lower epidermis is also smooth, with paracytic stomata;stomatal complex has sunken guard cells embedded below the overarching subsidiary cells with two cuticular ledges occurring between them;unicellular trichome bases can be found over the veins or in the areolas,surrounded by 5~6 radiating cells.It is more credible to decide the taxonomic position of fossil plant using leaf architecture and cuticular characteristics together.Compared with the modern leaf Litsea chunii,the fossil species has fewer trichome bases than the modern one.Consequently it was concluded that the increase of trichome bases resulted from their being in a cooler climate.Thus,based on the comparison in cuticular characteristics between the fossil and the modern leaves,it can be inferred that the climate was warmer and more humid in the pliocene of western Yunnan than the present.
A fossil leaf identified as Litsea cf.chunii Cheng was discovered in the late pliocene sediments of the Mangbang formation in Tengchong county,Yunnan province,China,which was well preserved with gross morphological structure and microstructure of cuticles in situ.The fossil leaf is narrowly elliptic,base cuneate, with obtuse apex and entire leaf margin.Its primary vein is straight and moderately thick;secondary veins are pinnate,which arise from the primary vein at angles from 30°~50°,and they are slightly curved near the leaf margin.Additionally,the anatomical characteristics of the present specimen are investigated as follows: the upper epidermis is free from stomata,composed of mostly tetragonal,pentagonal or slightly elongated cells,which are arranged in regular areola;anticlinal cell walls are straight and moderately thick,periclinal walls are smooth and slightly concave;trichome bases are rarely found.The lower epidermis is also smooth, with paracytic stomata;stomatal complex has sunken guard cells embedded below the overarching subsidiary cells with two cuticular ledges occurring between them;unicellular trichome bases can be found over the veins or in the areolas,surrounded by 5~6 radiating cells.It is more credible to decide the taxonomic position of fossil plant using leaf architecture and cuticular characteristics together.Compared with the modern leaf Litsea chunii,the fossil species has fewer trichome bases than the modern one.Consequently it was concluded that the increase of trichome bases resulted from their being in a cooler climate.Thus,based on the comparison in cuticular characteristics between the fossil and the modern leaves,it can be inferred that the climate was warmer and more humid in the pliocene of western Yunnan than the present.
引文
[1]中国科学院中国植物志编辑委员会.中国植物志(第三十一卷)[M].北京:科学出版社,1982:1-3; 261-339.
[2]DRINNAN A N,GRANE P R,FRIS E M, et al.Lauraceous flowers from the potomac group (mid-cretaceous) of eastern north America[J].Bot Gazette,1990,151(3):370-384.
[3]EKLUND H,KVACEK J.Lauraceous inflorescences and flowers from the cenomanian of Bohemia(Czech Republic,central Europe)[J].Int J Plant Sci,1998, 159(4):668-686.
[4]李锡文.樟科木姜子属群的起源与演化[J].云南植物研究,1995,17(3):251-254.
[5]WALKER J K.Comparative pollen morphology and phylogeny of the Ranalean complex[M]//BECK C B.Origin and Early Evolution of Angiosperms.New York:Columbia University Press,1976:241-299.
[6]HERENDEEN P S,CREPET W L,NIXON K C.Fossil flowers and pollen of Lauraceae from the upper cretaceous of New Jersey[J].P1 Syst and Evol,1994, 189(1/2):29-40.
[7]HU Ya-qin,KERGUSON D K,LI Cheng-sen,et al. Alseodaphne(Lauraceae) from the Pliocene of China and its paleoclimatic significance[J].Review of Palaeobotany and Palynology,2007,146(1/4): 277-285.
[8]DILCHER D L.Approaches to the identification of Angiosperm leaf remains[J].Bot Rev,1974,40(1): 1-157.
[9]KERP H.The study of fossil gymnosperms by means of cuticular analysis[J].Palaios,1990,5(6): 548-569.
[10]SUN Bai-nian,YAN De-fei,XIE San-ping,et al. Palaeogene fossil Populus leaves from Lanzhou basin and their palaeoclimatic significance[J].Chinese Science Bulletin,2004,49(13):1283-1289.
[11]HILL R S.Lauraceae leaves from the eocene of Nerriga,New South Wales[J].Alcheringa,1986, 10(4):327-351.
[12]Ow C,HSIAO J,LIAO C.Cuticle micromorphology of Cinnamomeae(Lauraceae) from Taiwan[J].Bull Exptl Forest NCHU,1992,14(2):1-30.
[13]BANDULSKA H.On the cuticles of some fossil and recent Lauraceae[J].Bot J Linn Soc,1926,47: 383-425.
[14]OHWI J.Flora of Japan[M].Washington D C: Smithsonian Institute,1965:110-116.
[15]KOSTERMANS A J G H.Lauraceae[J].Reinwardtia, 1957,4:193-256.
[16]李捷,李锡文.中国樟科木姜子属植物纪要[J].云南植物研究,2006,28(2):103-107.
[17]云南植被编写组.云南植被[M].北京:科学出版社, 1987:28-29.
[18]LIU Y S.Cuticular studies on two Pleistocene species of Lauraceae in Baise basin,Guangxi[J].Int J Plant Sci(Acta Bot Sin),1990,32(10):805-808.
[19]韦利杰,孙柏年,解三平,等.云南腾冲上新统植物油丹Alseodaphne hainanensis Merr.表皮微细构造研究[J].微体古生物学报,2005,22(4):392-399
[20]吴靖宇,孙柏年,解三平,等.云南腾冲新近系樟科润楠属两种化石及其古环境意义[J].高校地质学报,2008,14(1):90-98.
[21]朱华.云南种子植物区系地理成分分布格局及其意义[J].地球科学进展,2008,23(8):830-839.
[22]陶君容,周浙昆,刘裕生.中国晚白垩世至新生代植物区系发展演变[M].北京:科学出版社,2000: 73-77;92-103.
[23]姜汉侨.云南植被分布的特点及其地带规律性[J].云南植物研究,1980,2(1):22-31.
[24]戈宏儒,李代芸.云南西部新生代含煤盆地及聚煤规律[M].昆明:云南科技出版社,1999:20-23; 55-62.
[25]李锡康,谭筱虹,高子英,等.腾冲上新统芒棒组地质时代及沉积环境[J].云南地质,2004,23(2): 241-251.
[26]穆治国,佟伟,Garniss HC.腾冲火山活动的时代和岩浆来源问题[J].地球物理学报,1987(3):45-54.
[27]李大明,李齐,陈文寄.腾冲火山区上新世以来的火山活动[J].岩石学报,2000,16(3):362-370.
[28]CHRISTOPHEL D C,KERRIGAN R,ROWETT A I. The use of cuticular features in the taxonomy of the Lauraceae[J].Ann Mo Bot Gard,1996,83(3): 419-432.
[29]中国科学院北京植物研究所,南京地质古生物研究所《中国新生代植物》编写组.中国植物化石(第三册):中国新生代植物[M].北京:科学出版社, 1978:57-71.
[30]陶君荣,杜乃秋.云南腾冲新第三纪植物群及其时代[J].植物学报,1982,24(3):273-281. [31]KERP H.The study of fossil gymnosperms by means of cuticular analysis[J].Palaios,1990,5(6): 548-569.
[32]徐景先,王宇飞,杜乃秋.云南西部羊邑合龙林地区晚上新世植被和古气候[J].古地理学报,2003, 5(2):217-223.
[33]肖良,孙柏年,阎德飞,等.云南保山上新统黄背栎Quercus pannosa Hand.-Mazz.角质层特征及古环境意义[J].微体古生物学报,2006,23(1): 23-30.
[34]张绪教,何科昭,周志广.滇西地区新第三纪孢粉组合特征及环境变迁[J].现代地质,1996,10(2): 187-201.
[35]徐景先.云南中西部地区晚第三纪孢粉植物群及其古植被和古气候研究[D].北京:中国科学院植物研究所,2002.
[36]孙柏年,解三平,阎德飞,等.Ulmus harutoriensis Oishi et Huzioka角质层特征及古环境意义[J].兰州大学学报:自然科学版,2003,39(1):80-85.
[37]BANDULSKA H.A preliminary paper on the cuticular structure of certain dicotyledonous and coniferous leaves from the middle eocene flora of Bournemouth[J].Linn J Bot Soc,1923,46(308): 241-269.
[38]BANDULSKA H.On the cuticles of some recent and fossil Myrtaceae[J].Bot J Linn Soc,1931,48(325): 657-671.
[39]COLLINSON M E.Plant macrofossils from the Bracklesham group(early & middle) eocene,Bracklesham Bay,West Sussex,England:review and significance in the context of coeval British tertiary floras[J].Tert Res,1996,16(1/4):175-202.
[40]MCELWAIN J C.Do fossil plants signal palaeoat-mospheric CO_2 concentration in the geological past?[J].Phil Trans R Soc Lond B,1998,353(1365): 83-96.
[41]GREENWOOD D R,SCARR M J,CHRISTOPHEL D C.Leaf stomotal frequency in the Australian trop-ical rainforest tree Neolitsea dealbata(Lauraceae) as a proxy measure of atmospheric CO_2[J].Palaeo-geography, Palaeoclimatology,Palaeoecology, 2003,196(3/4):375-393.
[2]DRINNAN A N,GRANE P R,FRIS E M, et al.Lauraceous flowers from the potomac group (mid-cretaceous) of eastern north America[J].Bot Gazette,1990,151(3):370-384.
[3]EKLUND H,KVACEK J.Lauraceous inflorescences and flowers from the cenomanian of Bohemia(Czech Republic,central Europe)[J].Int J Plant Sci,1998, 159(4):668-686.
[4]李锡文.樟科木姜子属群的起源与演化[J].云南植物研究,1995,17(3):251-254.
[5]WALKER J K.Comparative pollen morphology and phylogeny of the Ranalean complex[M]//BECK C B.Origin and Early Evolution of Angiosperms.New York:Columbia University Press,1976:241-299.
[6]HERENDEEN P S,CREPET W L,NIXON K C.Fossil flowers and pollen of Lauraceae from the upper cretaceous of New Jersey[J].P1 Syst and Evol,1994, 189(1/2):29-40.
[7]HU Ya-qin,KERGUSON D K,LI Cheng-sen,et al. Alseodaphne(Lauraceae) from the Pliocene of China and its paleoclimatic significance[J].Review of Palaeobotany and Palynology,2007,146(1/4): 277-285.
[8]DILCHER D L.Approaches to the identification of Angiosperm leaf remains[J].Bot Rev,1974,40(1): 1-157.
[9]KERP H.The study of fossil gymnosperms by means of cuticular analysis[J].Palaios,1990,5(6): 548-569.
[10]SUN Bai-nian,YAN De-fei,XIE San-ping,et al. Palaeogene fossil Populus leaves from Lanzhou basin and their palaeoclimatic significance[J].Chinese Science Bulletin,2004,49(13):1283-1289.
[11]HILL R S.Lauraceae leaves from the eocene of Nerriga,New South Wales[J].Alcheringa,1986, 10(4):327-351.
[12]Ow C,HSIAO J,LIAO C.Cuticle micromorphology of Cinnamomeae(Lauraceae) from Taiwan[J].Bull Exptl Forest NCHU,1992,14(2):1-30.
[13]BANDULSKA H.On the cuticles of some fossil and recent Lauraceae[J].Bot J Linn Soc,1926,47: 383-425.
[14]OHWI J.Flora of Japan[M].Washington D C: Smithsonian Institute,1965:110-116.
[15]KOSTERMANS A J G H.Lauraceae[J].Reinwardtia, 1957,4:193-256.
[16]李捷,李锡文.中国樟科木姜子属植物纪要[J].云南植物研究,2006,28(2):103-107.
[17]云南植被编写组.云南植被[M].北京:科学出版社, 1987:28-29.
[18]LIU Y S.Cuticular studies on two Pleistocene species of Lauraceae in Baise basin,Guangxi[J].Int J Plant Sci(Acta Bot Sin),1990,32(10):805-808.
[19]韦利杰,孙柏年,解三平,等.云南腾冲上新统植物油丹Alseodaphne hainanensis Merr.表皮微细构造研究[J].微体古生物学报,2005,22(4):392-399
[20]吴靖宇,孙柏年,解三平,等.云南腾冲新近系樟科润楠属两种化石及其古环境意义[J].高校地质学报,2008,14(1):90-98.
[21]朱华.云南种子植物区系地理成分分布格局及其意义[J].地球科学进展,2008,23(8):830-839.
[22]陶君容,周浙昆,刘裕生.中国晚白垩世至新生代植物区系发展演变[M].北京:科学出版社,2000: 73-77;92-103.
[23]姜汉侨.云南植被分布的特点及其地带规律性[J].云南植物研究,1980,2(1):22-31.
[24]戈宏儒,李代芸.云南西部新生代含煤盆地及聚煤规律[M].昆明:云南科技出版社,1999:20-23; 55-62.
[25]李锡康,谭筱虹,高子英,等.腾冲上新统芒棒组地质时代及沉积环境[J].云南地质,2004,23(2): 241-251.
[26]穆治国,佟伟,Garniss HC.腾冲火山活动的时代和岩浆来源问题[J].地球物理学报,1987(3):45-54.
[27]李大明,李齐,陈文寄.腾冲火山区上新世以来的火山活动[J].岩石学报,2000,16(3):362-370.
[28]CHRISTOPHEL D C,KERRIGAN R,ROWETT A I. The use of cuticular features in the taxonomy of the Lauraceae[J].Ann Mo Bot Gard,1996,83(3): 419-432.
[29]中国科学院北京植物研究所,南京地质古生物研究所《中国新生代植物》编写组.中国植物化石(第三册):中国新生代植物[M].北京:科学出版社, 1978:57-71.
[30]陶君荣,杜乃秋.云南腾冲新第三纪植物群及其时代[J].植物学报,1982,24(3):273-281. [31]KERP H.The study of fossil gymnosperms by means of cuticular analysis[J].Palaios,1990,5(6): 548-569.
[32]徐景先,王宇飞,杜乃秋.云南西部羊邑合龙林地区晚上新世植被和古气候[J].古地理学报,2003, 5(2):217-223.
[33]肖良,孙柏年,阎德飞,等.云南保山上新统黄背栎Quercus pannosa Hand.-Mazz.角质层特征及古环境意义[J].微体古生物学报,2006,23(1): 23-30.
[34]张绪教,何科昭,周志广.滇西地区新第三纪孢粉组合特征及环境变迁[J].现代地质,1996,10(2): 187-201.
[35]徐景先.云南中西部地区晚第三纪孢粉植物群及其古植被和古气候研究[D].北京:中国科学院植物研究所,2002.
[36]孙柏年,解三平,阎德飞,等.Ulmus harutoriensis Oishi et Huzioka角质层特征及古环境意义[J].兰州大学学报:自然科学版,2003,39(1):80-85.
[37]BANDULSKA H.A preliminary paper on the cuticular structure of certain dicotyledonous and coniferous leaves from the middle eocene flora of Bournemouth[J].Linn J Bot Soc,1923,46(308): 241-269.
[38]BANDULSKA H.On the cuticles of some recent and fossil Myrtaceae[J].Bot J Linn Soc,1931,48(325): 657-671.
[39]COLLINSON M E.Plant macrofossils from the Bracklesham group(early & middle) eocene,Bracklesham Bay,West Sussex,England:review and significance in the context of coeval British tertiary floras[J].Tert Res,1996,16(1/4):175-202.
[40]MCELWAIN J C.Do fossil plants signal palaeoat-mospheric CO_2 concentration in the geological past?[J].Phil Trans R Soc Lond B,1998,353(1365): 83-96.
[41]GREENWOOD D R,SCARR M J,CHRISTOPHEL D C.Leaf stomotal frequency in the Australian trop-ical rainforest tree Neolitsea dealbata(Lauraceae) as a proxy measure of atmospheric CO_2[J].Palaeo-geography, Palaeoclimatology,Palaeoecology, 2003,196(3/4):375-393.
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