西藏芒康似勾儿茶叶属<bold>(</bold>鼠李科<bold>)</bold>化石及其生物地理学意义

周浙昆; 王腾翔; 黄健; 刘佳; 邓炜煜东; 李仕虎; 邓成龙; 苏涛

doi:10.1360/SSTe-2019-0099

SCIENTIA SINICA Terrae

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SCIENTIA SINICA Terrae, Volume 50, Issue 2: 233-244(2020) https://doi.org/10.1360/SSTe-2019-0099

西藏芒康似勾儿茶叶属(鼠李科)化石及其生物地理学意义

周浙昆^1,2,*,†, 王腾翔^1,3,†, 黄健¹, 刘佳¹, 邓炜煜东^1,3, 李仕虎^4,5, 邓成龙⁴, 苏涛^1,3,§

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ReceivedMay 7, 2019
AcceptedAug 23, 2019
PublishedNov 26, 2019

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Abstract

文章报道了发现于西藏东南部芒康盆地上始新统的鼠李科植物叶化石标本. 化石叶缘全缘或微波状, 二级脉间距规则, 弧曲向上并与其上的二级脉在叶缘处结合并与叶缘融合, 形成真曲脉序; 二级脉在中上部较为密集, 三级脉密集相互平行, 横贯二级脉之间, 这些叶脉特征与美洲分布的灭绝类群——似勾儿茶叶属(Berhamniphyllum Jones and Dilcher)非常相似. 百分之四十的二级脉集中于叶的上半部是当前化石的一个典型特征, 未见于该属的其他类群, 于是将当前化石定为君容似勾儿茶叶新种(Berhamniphyllum junrongiae Z. K. Zhou, T. X. Wang et J. Huang sp. nov.). 进一步研究发现, 仅凭叶脉特征不能将勾儿茶属(Berchemia)和Rhamnidium、Karwinskia等三个属区分开来, 似勾儿茶叶属(Berhamniphyllum)代表了这几个属的一个灭绝的共同祖先, 本文对产于云南和山东等地三种勾儿茶属化石进行了归并, 并将其归并入似勾儿茶叶属中. 根据形态学、分子系统证据和化石记录, 将勾儿茶属、Rhamnidium、Karwinskia和似勾儿茶叶属定义为勾儿茶复合群(Berchemia complex). 本文还讨论了勾儿茶复合群地理分布格局的演变历史, 认为勾儿茶复合群于晚白垩世晚期起源于南美哥伦比亚, 在始新世经中美洲扩散到北美, 后又从北美经过北大西洋陆桥扩散至欧洲并从欧洲扩散至非洲. 东亚的勾儿茶复合群是经白令陆桥扩散而来, 时间不晚于始新世, 这一类群在东亚最早出现于西藏芒康, 其后再扩散至亚洲其他地区.

Funded by

中国科学院战略性先导科技专项项目(XDA2007030102,XDB26000000)

国家自然科学基金-英国自然环境研究理事会国际合作重点项目(41661134049,NE/P013805/1)

中国科学院第二次青藏高原综合科学考察研究项目(STEP)

中国科学院青年创新促进会项目(2017439)

中国科学院前沿科学重点研究计划项目(QYZDB-SSW-SMC016)

Acknowledgment

感谢中国科学院西双版纳热带植物园和昆明植物研究所的同事参与野外工作. 感谢西藏芒康卡均藏族同胞在野外工作中给予的帮助. 感谢史恭乐博士帮助拍摄云南开远小龙潭组勾儿茶的化石照片, Lutz Kunzmann教授、王雨晴和Steven Manchester教授提供了部分欧洲、日本和北美的文献. 在论文写作过程中, 与贾林波博士和Steven Manchester教授进行过有益的讨论. 感谢中国科学院西双版纳热带植物园中心实验室在摄影设备和技术上提供的支持. 感谢两位匿名审稿人在审稿过程中提出意见和建议, 这些意见和建议对于拙稿的改进起到了重要作用.

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图 1
勾儿茶属及其相似类群的化石记录和现代分布
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图 2
化石采集点位置
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图 3
君容似勾儿茶叶(新种)(Berhamniphyllum junrongiae Z. K. Zhou, T.X. Wang et J. Huang sp. nov.)、多花似勾儿茶叶(Berhamniphyllum miofloribundum (Hu et Chaney) J. Huang, T. Su et Z. K. Zhou comb. nov.)及长梗勾儿茶(Berchemia longipes)(现代)透明叶
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图 4
勾儿茶属、Karwinskia属及Rhamnidium属
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图 5
鼠李族的勾儿茶类植物可能的传播路径

表 1 勾儿茶属化石记录

物种	器官	时代	地点	文献
Berchemia mellerae	内果皮	中始新世	德国Messel	Collinson等, 2012
B. sp	叶	晚始新世	美国俄勒冈州	Manchester, 2000
B. altorhenana	叶	早渐新世	德国Rauenberg	Kovar-Eder, 2016
B. multinervis	叶	晚渐新世	瑞士Ebnat-Kappel	Büchler, 1990
B. multinervis	叶	晚渐新世	保加利亚Vulche Pole Molasse组	Bozukov等, 2008
B. multinervis	叶	晚渐新世	罗马尼亚Petrosani盆地	Givulescu, 1996
B. huanoides	叶	晚渐新世	美国蒙大拿州	Becker, 1969
B. multinervis	叶	中新世	保加利亚Western Rhodopes	Bozukov, 2000
B. priscaformis	叶	中新世	美国南卡罗莱纳州	Berry, 1916b
B. sp	叶	中新世	美国爱达荷州	Smiley等, 1975
B. pseudodiscolor	种子	早中新世	肯尼亚Rusinga岛	Collinson等, 2009
B. acutangula	叶	早中新世	巴伐利亚Goldern	Spitzelberger, 1989
B. miofloribunda	叶	早中新世	日本本州	Yabe, 2008
B. multinervis	叶	早中新世	塞尔维亚Valjevo-Mionica盆地	Lazarević等, 2013
B. multinervis	叶	早中新世	捷克Most盆地	Sakala, 2000; Teodoridis, 2007
B. multinervis	叶	早-中中新世	瑞士Canton Lucerne	Köecke和Uhl, 2015
B. pseudodiscolor	外果皮	中中新世	肯尼亚Fort Ternan	Retallack, 1992
B. miofloribunda	叶	中中新世	日本能登半岛	Ishida, 1970
B. nepalensis	叶	中中新世	尼泊尔Koilabas	Prasad和Dwivedi, 2007
B. multinervis	叶	晚中新世	巴伐利亚Lerch	Jung, 1968
B. multinervis	叶	晚中新世	瑞士Oehingen	Hantke, 1954; Heer, 1855–1859
B. miofloribunda	叶	晚中新世	日本本州	Ozaki, 1980
B. miofloribunda	叶	晚中新世	日本本州	Ozaki, 1991
B. miofloribunda	叶	上新世	中国云南团田	吴靖宇, 2009
B. cf. yunnanensis	叶	上新世	中国云南团田	吴靖宇, 2009
B. multinervis	叶	上新世	意大利Fossano	Macaluso等, 2018
B. floribunda	叶	晚第三纪	印度Jharkhand	Singh和Prasad, 2007

表 2 其他勾儿茶类化石记录

物种	时代	地点	文献
Berhamniphyllum sp.	晚白垩世	南美哥伦比亚	Correa等, 2010
Berhamniphyllum claibornense	早始新世	美国肯塔基州、田纳西州	Berry, 1916a; Jones和Dilcher, 1980
Berhamniphyllum claibornense	中始新世	美国田纳西州	Dilcher和Lott, 2005
Berhamniphyllum sp.	中始新世	美国田纳西州	Dilcher和Lott, 2005
Berhamniphyllum sp.	晚始新世	美国俄勒冈州	Myers等, 2002
Berhamniphyllum junrongiae sp. nov.	晚始新世	中国西藏芒康	本研究
Karwinskia axamilpense	渐新世	墨西哥普埃布拉	de León等, 1998
Berhamniphyllum miofloribundum comb. nov.	中新世	中国山东山旺; 云南文山、小龙潭	中国新生代植物编写组, 1978; 周浙昆, 1985; 黄健, 2017; 本研究

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西藏芒康似勾儿茶叶属(鼠李科)化石及其生物地理学意义

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