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SCIENCE CHINA Earth Sciences
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SCIENCE CHINA Earth Sciences, Volume 63, Issue 2: 224-234(2020) https://doi.org/10.1007/s11430-019-9477-8

Fossil leaves of Berhamniphyllum (Rhamnaceae) from Markam, Tibet and their biogeographic implications

Zhekun ZHOU1,2,*,†, Tengxiang WANG1,3,†, Jian HUANG1, Jia LIU1, Weiyudong DENG1,3, Shihu LI4,5, Chenglong DENG4, Tao SU1,3,§
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  • ReceivedMay 7, 2019
  • AcceptedAug 23, 2019
  • PublishedNov 22, 2019
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Abstract

A new occurrence of buckthorn fossil leaves is reported from the upper Eocene strata of Markam Basin, southeastern Tibet, China. The leaf margin is either entire or slightly sinuous. Secondary veins are regularly spaced, forming eucamptodromous venation. These secondaries exist as straight lines from midvein to near margin and then arch abruptly upward and enter into a margin vein. The tertiary veins are densely spaced and parallel, and are percurrent to secondary veins. This leaf architecture conforms with Berhamniphyllum Jones and Dilcher, an extinct fossil genus reported from America. Our fossils are characterized by their dense secondaries, with secondary veins on the upper half portion of the blade accounting for over 40% of all secondaries. A new species, Berhamniphyllum junrongiae Z. K. Zhou, T. X. Wang et J. Huang sp. nov., is proposed. Further analysis shows that confident assignment among Rhamnidium, Berchemia, and Karwinskia cannot be made based on leaf characters alone. Berhamniphyllum might represent an extinct common ancestor of these genera. In this study, several fossil Berchemia from Yunnan and Shandong are emended and reassigned to Berhamniphyllum. A new complex, namely the Berchemia Complex, is proposed based on morphology, molecular evidence, and the fossil record. This complex contains the fossil leaves of Rhamnidium, Karwinskia, Berchemia, and Berhamniphyllum. The historical biogeography of the Berchemia Complex is also discussed in this paper. This complex might have originated in the late Cretaceous in Colombia, South America, and dispersed to North America via Central America during the Eocene. Subsequently, the complex moved from North America to East Asia via the Bering Land Bridge no later than the late Eocene. Besides, the complex migrated from North America to Europe via the North Atlantic Land Bridge and then migrated further to Africa. In East Asia, it first appeared in Markam on the Qinghai-Tibetan Plateau, and then dispersed to other regions of Asia.


Funded by

the Strategic Priority Research Program of CAS(Grant,Nos.,XDA2007030102,&,XDB26000000)

the NSFC(the,National,Natural,Science,Foundation,of,China)

CAS(Grant,No.,2019QZKK0705)

Youth Innovation Promotion Association

CAS(Grant,No.,2017439)

Key Research Program of Frontier Sciences

CAS(Grant,No.,QYZDB-SSW-SMC016)


Acknowledgment

We thank colleagues from Xishuangbanna Tropical Botanical Garden (XTBG), Chinese Academy of Sciences (CAS), and Kunming Institute of Botany, CAS for field work; Tibetan villagers from Kajun village for their kind help during field work; Dr. Gongle Shi for photographing fossil from Xiaolongtan flora; Prof. Lutz Kunzmann, Ms. Yuqing Wang and Prof. Steven Manchester for providing literature; Dr. Linbo Jia and Prof. Steven Manchester for discussion and comments; the Public Technology Service Center, XTBG, CAS for providing microscopes and experimental facilities; Teresa Spicer for improving the English manuscript. We are also grateful to two anonymous reviewers for their constructive advices. This study was supported by the Strategic Priority Research Program of CAS (Grant Nos. XDA2007030102 & XDB26000000), the NSFC (the National Natural Science Foundation of China)-NERC (Natural Environment Research Council of the United Kingdom) joint research program (Grant Nos. 41661134049 & NE/P013805/1); The Second Tibetan Plateau Scientific Expedition and Research Program (STEP), CAS (Grant No. 2019QZKK0705), Youth Innovation Promotion Association, CAS (Grant No. 2017439) and Key Research Program of Frontier Sciences, CAS (Grant No. QYZDB-SSW-SMC016).


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  • Click to view Download high-quality image

    Figure 1

    Historical and modern distribution of Berchemia and resembling taxa.

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    Figure 2

    Fossil locality in Yunnan and Tibet, China.

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    Figure 3

    Berhamniphyllum fossils reported in this study and cleared leaf of extant species Berchemia longipes. (a)–(c) Berhamniphyllum junrongiae Z. K. Zhou, T. X. Wang et J. Huang sp. nov., (a) MK3-1491, scale bar=1 cm; (b) Details of MK3-1491, S, secondary veins; IS, inter secondary veins; PT, percurrent tertiary veins; DT, dichotomous tertiary veins, scale bar=0.5 cm; (c) MK3-2155, scale bar=0.5 cm. (d)–(f) Berhamniphyllum miofloribundum (Hu et Chaney) J. Huang, T. Su et Z. K. Zhou comb. nov., (d) DMS-1713, scale bar=0.5 cm; (e) DMS-3872, scale bar=0.5 cm; (f) PB12519, scale bar=0.5 cm. (g) Line drawing of MK3-1491, scale bar=0.5 cm; (h) Cleared leaf of Berchemia longipes, scale bar=1 cm.

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    Figure 4

    Specimens of Berchemia, Karwinsia and Rhamnidium. (a) Berchemia longipes, photograghed from Xishuangbanna Tropical Botanical Garden, CAS, leaves alternate; (b) Karwinskia calderonii, scale bar=5 cm, extracted from JSTOR, leaves opposite; (c) Rhamnidium caloneurum, scale bar=5 cm, extracted from JSTOR, leaves opposite.

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    Figure 5

    Hypothesized migration route modified from Jia et al. (2015).

  • Table 1   Fossil record of

    Species

    Organ

    Age

    Locality

    Reference

    Berchemia mellerae

    Endocarp

    Middle Eocene

    Messel, Germany

    Collinson et al., 2012

    B. sp

    Leaf

    Late Eocene

    Wheeler County, Oregon, USA

    Manchester, 2000

    B. altorhenana

    Leaf

    Early Oligocene

    Rauenberg, Germany

    Kovar-Eder, 2016

    B. multinervis

    Leaf

    Late Oligocene

    Ebnat-Kappel, Switzerland

    Büchler, 1990

    B. multinervis

    Leaf

    Late Oligocene

    Vulche Pole Molasse Formation, SE Bulgaria

    Bozukov et al., 2008

    B. multinervis

    Leaf

    Late Oligocene

    Petrosani Basin, Romania

    Givulescu, 1996

    B. huanoides

    Leaf

    Late Oligocene

    Beaverhead County, Montana, USA

    Becker, 1969

    B. multinervis

    Leaf

    Miocene

    Western Rhodopes, Bulgaria

    Bozukov, 2000

    B. priscaformis

    Leaf

    Miocene

    South Caroline, USA

    Berry, 1916b

    B. sp

    Leaf

    Miocene

    Clarkia, Idaho, USA

    Smiley et al., 1975

    B. pseudodiscolor

    Seed

    Early Miocene

    Rusinga Island, Kenya

    Collinson et al., 2009

    B. acutangula

    Leaf

    Early Miocene

    Goldern, Bavaria

    Spitzelberger, 1989

    B. miofloribunda

    Leaf

    Early Miocene

    Honshu, Japan

    Yabe, 2008

    B. multinervis

    Leaf

    Early Miocene

    Valjevo-Mionica Basin, Serbia

    Lazarević et al., 2013

    B. multinervis

    Leaf

    Early Miocene

    Most Basin, Czech

    Sakala, 2000; Teodoridis, 2007

    B. multinervis

    Leaf

    Early-middle Miocene

    Canton Lucerne, Switzerland

    Köecke and Uhl, 2015

    B. pseudodiscolor

    exocarp

    Middle Miocene

    Fort Ternan, Kenya

    Retallack, 1992

    B. miofloribunda

    Leaf

    Middle Miocene

    Noto Peninsula, Japan

    Ishida, 1970

    B. nepalensis

    Leaf

    Middle Miocene

    Koilabas, Nepal

    Prasad and Dwivedi, 2007

    B. multinervis

    Leaf

    Late Miocene

    Lerch, Bavaria

    Jung, 1968

    B. multinervis

    Leaf

    Late Miocene

    Oehingen, Switzerland

    Hantke, 1954; Heer, 1855–1859

    B. miofloribunda

    Leaf

    Late Miocene

    Honshu, Japan

    Ozaki, 1980

    B. miofloribunda

    Leaf

    Late Miocene

    Honshu, Japan

    Ozaki, 1991

    B. calymmatophyllua

    Leaf

    Late Miocene

    Lincang, Yunnan Province, China

    Guo, 2011

    B. miofloribunda

    Leaf

    Pliocene

    Tuantian, Yunnan Province, China

    Wu, 2009

    B. cf. yunnanensis

    Leaf

    Pliocene

    Tuantian, Yunnan Province, China

    Wu, 2009

    B. multinervis

    Leaf

    Pliocene

    Fossano, Italy

    Macaluso et al., 2018

    B. floribunda

    Leaf

    LateTertiary

    Jharkhand, India

    Singh and Prasad, 2007

  • Table 2   Fossil records of affinities

    Species

    Age

    Locality

    Reference

    Berhamniphyllum sp.

    Late Cretaceous

    Paz de Río, Boyacá, Columbia

    Correa et al., 2010

    Berhamniphyllum claibornense

    Early Eocene

    Puryear County and Henry County, Tennessee;Graves County, Kentucky

    Berry, 1916a; Jones and Dilcher, 1980

    Berhamniphyllum claibornense

    Middle Eocene

    Powers Pit, Western Tennessee

    Dilcher and Lott, 2005

    Berhamniphyllum sp.

    Middle Eocene

    Powers Pit, Western Tennessee

    Dilcher and Lott, 2005

    Berhamniphyllum sp.

    Late Eocene

    The Willamette flora, Oregon

    Myers et al., 2002

    Berhamniphyllum junrongiae

    Late Eocene

    Markam, Tibet, China

    This study

    Karwinskia axamilpense

    Oligocene

    Puebla, Mexico

    de León et al., 1998

    Berhamniphyllum miofloribundum

    Miocene

    Shanwang, Shandong Province, China; Wenshanand Xiaolongtan, Yunnan Province, China

    WGCPC, 1978; Zhou, 1985;Huang, 2017; this study

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