云南保山上新统前灰背栎Quercus presenescens角质层特征及古气候意义
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摘要
在云南保山羊邑煤矿上新统羊邑组发现了一栎属高山栎组植物压型化石,根据叶形态和表皮微细构造特征,将其鉴定为前灰背栎(Quercus presenescens Z.K.Zhou)。该标本具有之字形中脉,叶最宽处形成一矩形区域,叶缘略反卷的特征,与已报道的前灰背栎化石形态一致,且区别于其它已发现的高山栎组化石。解剖学研究表明,该化石叶片为气孔下生型,上、下表皮均无毛基;上表皮细胞为四边形,垂周壁波状弯曲;下表皮细胞多为四-五边形,垂周壁波状弯曲,气孔器环列型。通过与现生高山栎组植物叶形态相比,当前化石与灰背栎最为接近,且其表皮细胞形态、气孔器类型与现生灰背栎基本一致,二者的主要区别在于前灰背栎无毛基,而现生灰背栎有表皮毛。研究认为,两者被毛的差异,是植物叶形态结构对气候变化的响应。
A compression fossil reported as Quercus sect.Heterobalanus was collected from the Pliocene Yangyi Formation at the Yangyi coal-mine of Bao-shan,Yunnan,and is identified in this study as Quercus presenescens Z.K.Zhou based on morphological and cuticular characters.The typical morphological characters of the fossil are zigzag primary vein,oblong leaf blade and revolute leaf margin;these features are characteristic of Q.pre-senescens,but different from those of Quercus sect.Heterobalanus.The epidermal characters are as follows:stomata distributing on the lower epidermis,no trichomes on the upper and lower epidermis;cells on the upper epidermis quadrangular,anticlinal cell walls undulate;cells on the lower epidermis quadrangular to pentagonal,anticlinal cells wall undulate,stomata cyclocytic.The specimen shows closer affinities with the living species Q.senescens based on a detailed comparison of leaf architecture with living species of Quercus sect.Heterobalanus.Also the cell shape and the type of stomata are identical to these of the living species Q.senescens.The fossil is different from living Q.senescens in having no trichome.This implies that the common existence of trichomes in living Quercus sect.Heterobalanus may have been the result of adaptation to recent climatic change.
A compression fossil reported as Quercus sect.Heterobalanus was collected from the Pliocene Yangyi Formation at the Yangyi coal-mine of Bao-shan,Yunnan,and is identified in this study as Quercus presenescens Z.K.Zhou based on morphological and cuticular characters.The typical morphological characters of the fossil are zigzag primary vein,oblong leaf blade and revolute leaf margin;these features are characteristic of Q.pre-senescens,but different from those of Quercus sect.Heterobalanus.The epidermal characters are as follows:stomata distributing on the lower epidermis,no trichomes on the upper and lower epidermis;cells on the upper epidermis quadrangular,anticlinal cell walls undulate;cells on the lower epidermis quadrangular to pentagonal,anticlinal cells wall undulate,stomata cyclocytic.The specimen shows closer affinities with the living species Q.senescens based on a detailed comparison of leaf architecture with living species of Quercus sect.Heterobalanus.Also the cell shape and the type of stomata are identical to these of the living species Q.senescens.The fossil is different from living Q.senescens in having no trichome.This implies that the common existence of trichomes in living Quercus sect.Heterobalanus may have been the result of adaptation to recent climatic change.
引文
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Leng Qin,2000b.Fagaceae fromthe Miocene Daotaiqiao Formationof Huanan,Heilongjiang,China.Acta Palaeontologica Sinica(古生物学报),39(Suppl.):251—287.
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Tang Hua-fen(唐华芬),Hu You-heng(胡友恒),1993.Late Pliocenepalynolflora of bearing coal sediments in western Yunnan.CoalGeology China(中国煤炭地质),5(4):25—29(in Chinese).
Tao Jun-rong(陶君容),Zhou Zhe-kun(周浙昆),Liu Yu-sheng(刘裕生),2000.The Evolution of the Late Cretaceous-Cenozoic Flo-ras in China.Beijing:Science Press.1—282(in Chinese).
The Writing Group of Cenozoic Plant of China(中国新生代植物编写组),1978.Fossil Plants of China,Cenozoic Plants from China,3.Beijing:Science Press.1—232(in Chinese).
Uzunova K,Palamarev E,Ehrendorfer F,1997.Anatomical changesand evolutionary trendsinthe foliar epidermis of extant andfos-sil Euro-Mediterranean oaks(Fagaceae).Plant Systematics andEvolution,204:141—159.
Xiao Liang(肖良),Sun Bai-nian(孙柏年),Yan De-fei(阎德飞)etal.,2006.Cuticular structure ofQuercus pannosaHand.-Mazz.fromthe Pliocene in Baoshan,Yunnan Province and its palaeo-environmental significance.Acta Micropalaeontologica Sinica(微体古生物学报),23(1):23—30(in Chinese with English ab-stract).
Xu Jing-xian(徐景先),Wang Yu-fei(王宇飞),Du Nai-qiu(杜乃秋),2003.Late Pliocene vegetation and palaeoclimate of Yangyi andLongling of western Yunnan Province.Journal of Palaeogeogra-phy(古地理学报),5(2):217—223(in Chinese with English ab-stract).
Xu Jing-xian,Blackmore S,Wang Yu-feiet al.,2004.Late Pliocenevegetation and climate of Yangyi region Yunnan of China,basedon palynological data.Palaeontographica Abteilung B,269:131—148.
Xu Ren(徐仁),Tao Jun-rong(陶君容),Sun Xiang-jun(孙湘君),1973.On the discovery of aQuercus semicarpifoliabed inmount Shiha Pangma andits significancein botany and geology.Acta Botanica Sinica(植物学报),15(1):103—114(in Chinese).
Xu Yong-chun(徐永椿),Ren Xian-qi(任宪戚),1976.Class and dis-tribution of Fagaceae in Yunnan(Ⅱ),Acta PhytotaxonomicaSinica(植物分类学报),14(2):79—86(in Chinese).
Yun Bo(云博),1975.New material of about elephant fossil fromYunnan Province.Vertebrata Palaxiatica(古脊椎动物学报),13(4):229—233(in Chinese).
Zhou Zhe-kun(周浙昆),1992.Ataxonomical revision of fossil ever-green sclerophyllous oaks from China.Acta Botanica Sinica(植物学报),34(2):954—961(in Chinese).
Zhou Zhe-kun(周浙昆),1993.Geographical distribution ofQuercusfrom China.Journal of Graduate School Academia Sinica(中国科学院研究生院学报),10(1):95—102(in Chinese with Englishabstract).
Zhou Zhe-kun(周浙昆),Pu Chun-xia(普春霞),Chen Wen-yun(陈文允),2003.Relationships between the distributions ofQuercussect.Heterobalanus(Fagaceae)and uplift of Himalayas.Ad-vance in Earth Sciences(地球科学进展),18(6):884—890(inChinese with English abstract).
Zhou Zhe-Kun(周浙昆),Yang Qing-song(杨青松),Xia Ke(夏珂),2007.Fossils ofQuercussect.Heterobalanuscan help ex-plain the uplift of the Himalayas.Chinese Science Bulletin(科学通报),52(2):238—247(Chinese edition).
Zhou Zhe-kun,Wilkinson HP,Wu Zheng-yi,1995.Taxonomical andevolutionaryimplications of the leaf anatomy and architecture ofQuercusL.subgenusQuercusfrom China.Cathaya,7:1—34.
Dilcher D L,1974.The botanical review(Interpreting Botanical Pro-gress).The New York Botanical Garden,40(1):1—141.
Ge Hong-ru(戈宏儒),Li Dai-yun(李代芸),1999.Cenozoic Coal-bearing Basins and Coal-forming Regularity in West Yunnan.Kunming:Yunnan Science and Technology Press.1—88(in Chi-nese).
He Jin-sheng(贺金生),Chen Wei-lie(陈伟烈),Wang Xun-ling(王勋陵),1994.Morphological and anatomical features ofQuercussectionSuberand its adaptation to the ecological environment.Acta Phytoecologica Sinica(植物分类学报),18(3):219—227(inChinese).
Kerp H,1990.The study of fossil Gymnosperms by means of cuticu-lar analysis.Palaios,5:548—569.
Leng Qin,2000a.Arevision ofQuercus miocrispularHuzioka(sub-sect.Diversipilosae,sect.Lepidobalanus)fromthe Miocene ofeastern Asia.Paleontological Journal,34(3):341—348.
Leng Qin,2000b.Fagaceae fromthe Miocene Daotaiqiao Formationof Huanan,Heilongjiang,China.Acta Palaeontologica Sinica(古生物学报),39(Suppl.):251—287.
Li Hao-min(李浩敏),Guo Shuang-xing(郭双兴),1976.The Mio-cene flora Namling of Xizang.Acta Palaeontologica Sinica(古生物学报),15(1):7—17(in Chinese with English abstract).
Li Hao-min,Hickey L J,1988.Leaf architecture and systematic ofthe Hamamelidaceae sensu lato.Acta Phytotaxonomica Sinica(植物分类学报),26(2):96—110.
Liu Yu-sheng(刘裕生),Zheng Ya-hui(郑亚惠),1995.Neogene flo-ras.In:Li Xing-xue(李星学)(ed.),Fossil Floras of Chinathrough the Geological Ages.Guangzhou:Guangdong Scienceand Technology Press.383—416(in Chinese).
Luo Yan,Zhou Zhe-kun,2002.Leaf architecture inQuercussubge-nusCyclobalanopsis(Fagaceae)from China.Botanical Journalof the Linnean Society,140:283—295.
Pan Kai-yu(潘开玉),Lu An-ming(路安民),Wen Jie(温洁),1990.Characters of leaf epidermis in Hamamelidaceae(S.L.).Acta Phytotaxonomica Sinica(植物分类学报),28(1):10—26(inChinese with English summary).
Sun Bai-nian(孙柏年),Cong Pei-yun(丛培允),Yan De-fei(阎德飞)et al.,2003.Cuticular structure of two angiosperm fossils inNeogene from Tengchong,Yunnan Province and its palaeoenvi-ronmental significance.Acta Palaeontologica Sinica(古生物学报),42(2):216—221(in Chinese with English abstract).
Sun Bai-nian(孙柏年),Yan De-fei(阎德飞),Xie San-ping(解三平)etal.,2004.Palaeogene fossilPopulusleaves fromLanzhou Basinandtheir palaeoclimate significance.Chinese Science Bulletin(科学通报),2004,49(13):1283—1289(in Chinese with Englishabstract).
Sun Hang(孙航),Chen Jie(陈介),Zhou Zhe-kun(周浙昆)etal.,1991.Theleaf architecture andits taxonomic significanceinthe generaAlbiziaandCylindrokeluphafrom China.Acta Bo-tanica Yunnanica(云南植物研究),13(3):241—253(in Chinesewith English abstract).
Sun Hang(孙航),2002.Tethys retreat and Himalayas-Hengduan-shan mountains uplift andtheir significance onthe origin and de-velopment of the Sino-Himalayan elements and alpine flora.Ac-ta Botanica Yunnanica(云南植物研究),24(3):273—288(inChinese with English abstract).
Tang Hua-fen(唐华芬),Hu You-heng(胡友恒),1993.Late Pliocenepalynolflora of bearing coal sediments in western Yunnan.CoalGeology China(中国煤炭地质),5(4):25—29(in Chinese).
Tao Jun-rong(陶君容),Zhou Zhe-kun(周浙昆),Liu Yu-sheng(刘裕生),2000.The Evolution of the Late Cretaceous-Cenozoic Flo-ras in China.Beijing:Science Press.1—282(in Chinese).
The Writing Group of Cenozoic Plant of China(中国新生代植物编写组),1978.Fossil Plants of China,Cenozoic Plants from China,3.Beijing:Science Press.1—232(in Chinese).
Uzunova K,Palamarev E,Ehrendorfer F,1997.Anatomical changesand evolutionary trendsinthe foliar epidermis of extant andfos-sil Euro-Mediterranean oaks(Fagaceae).Plant Systematics andEvolution,204:141—159.
Xiao Liang(肖良),Sun Bai-nian(孙柏年),Yan De-fei(阎德飞)etal.,2006.Cuticular structure ofQuercus pannosaHand.-Mazz.fromthe Pliocene in Baoshan,Yunnan Province and its palaeo-environmental significance.Acta Micropalaeontologica Sinica(微体古生物学报),23(1):23—30(in Chinese with English ab-stract).
Xu Jing-xian(徐景先),Wang Yu-fei(王宇飞),Du Nai-qiu(杜乃秋),2003.Late Pliocene vegetation and palaeoclimate of Yangyi andLongling of western Yunnan Province.Journal of Palaeogeogra-phy(古地理学报),5(2):217—223(in Chinese with English ab-stract).
Xu Jing-xian,Blackmore S,Wang Yu-feiet al.,2004.Late Pliocenevegetation and climate of Yangyi region Yunnan of China,basedon palynological data.Palaeontographica Abteilung B,269:131—148.
Xu Ren(徐仁),Tao Jun-rong(陶君容),Sun Xiang-jun(孙湘君),1973.On the discovery of aQuercus semicarpifoliabed inmount Shiha Pangma andits significancein botany and geology.Acta Botanica Sinica(植物学报),15(1):103—114(in Chinese).
Xu Yong-chun(徐永椿),Ren Xian-qi(任宪戚),1976.Class and dis-tribution of Fagaceae in Yunnan(Ⅱ),Acta PhytotaxonomicaSinica(植物分类学报),14(2):79—86(in Chinese).
Yun Bo(云博),1975.New material of about elephant fossil fromYunnan Province.Vertebrata Palaxiatica(古脊椎动物学报),13(4):229—233(in Chinese).
Zhou Zhe-kun(周浙昆),1992.Ataxonomical revision of fossil ever-green sclerophyllous oaks from China.Acta Botanica Sinica(植物学报),34(2):954—961(in Chinese).
Zhou Zhe-kun(周浙昆),1993.Geographical distribution ofQuercusfrom China.Journal of Graduate School Academia Sinica(中国科学院研究生院学报),10(1):95—102(in Chinese with Englishabstract).
Zhou Zhe-kun(周浙昆),Pu Chun-xia(普春霞),Chen Wen-yun(陈文允),2003.Relationships between the distributions ofQuercussect.Heterobalanus(Fagaceae)and uplift of Himalayas.Ad-vance in Earth Sciences(地球科学进展),18(6):884—890(inChinese with English abstract).
Zhou Zhe-Kun(周浙昆),Yang Qing-song(杨青松),Xia Ke(夏珂),2007.Fossils ofQuercussect.Heterobalanuscan help ex-plain the uplift of the Himalayas.Chinese Science Bulletin(科学通报),52(2):238—247(Chinese edition).
Zhou Zhe-kun,Wilkinson HP,Wu Zheng-yi,1995.Taxonomical andevolutionaryimplications of the leaf anatomy and architecture ofQuercusL.subgenusQuercusfrom China.Cathaya,7:1—34.
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