Research papers
Fokienia shengxianensis sp. nov. (Cupressaceae) from the late Miocene of eastern China and its paleoecological implications

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Abstract

Twenty-one fossil foliage, identified as Fokienia shengxianensis sp. nov. (Cupressaceae), were collected from the upper Miocene Shengxian Formation in Tiantai and Ninghai counties, Zhejiang Province of eastern China. These fossils can likely be distinguished from other extant genera in the family Cupressaceae s.l. except Fokienia by leaf external morphology alone. Foliar cuticular comparison with the only modern species in this genus, Fokienia hodginsii Henry et Thomas, further suggests that the present fossil species closely resembles the modern counterpart. Paleoecologically, the occurrence of Fokienia from the Shengxian Formation appears to support that the fossil site was covered by subtropical evergreen broad-leaved forest and under warm and humid conditions during the late Miocene, consistent with evidence from previously published fossil plants.

Highlights

► Miocene Fokienia leaves from China ► First fossil record of Fokienia with cuticle ► Support warm and humid environment in eastern China during the Miocene

Introduction

Fokienia, one of the 30 genera in the Cupressaceae s.l., is a monotypic genus with only one modern species, Fokienia hodginsii (Farjon, 2005). Today, Fokienia occurs in southern China, northern Vietnam, and northern Laos (Zheng and Fu, 1978). This conifer is usually a minor constituent of a subtropical evergreen (mixed) mesophytic forest, which in an undisturbed state is dominated by numerous angiospermous trees, e.g., Quercus spp., Lithocarpus spp., Pasania spp., Nyssa sinensis, Schima argentea, Schima superba, and a few other conifers, e.g., Cephalotaxus fortunei and Nothotsuga longibracteata (Farjon, 2005). It grows in mountains at about 100–1800 m above sea level (Zheng and Fu, 1978) where the mean annual precipitation is 1200 mm or more (Farjon, 2005).

Fokienia is very rare in the fossil record. McIver and Basinger (1990) reported fossil Fokienia foliage and attached seed cones from the Paleocene of Saskatchewan, central Canada. However, the taxonomic treatment of this fossil species has been questioned by Manchester et al. (2009), who suggested that it may belong to Ditaxocladus, an extinct cupressaceous genus. Another fossil record, represented by foliage twigs with two nodes, was reported from the Oligocene of Jilin, northeastern China (Guo and Zhang, 2002). Generally, due to the great similarity of foliar characters in the Cupressaceae, their fossil identifications are best based on seed cone structure (Offler, 1984, McIver and Basinger, 1987, McIver, 1989, McIver and Basinger, 1990). Unfortunately, cupressaceous remains usually consist of foliage only without attached seed cones (e.g.,Chaney and Axelrod, 1959, Brown, 1962, Li and Guo, 1982, Guo and Zhang, 2002). However, recent work has suggested that cuticular characters of cupressaceous fossil leaves can be diagnostic (e.g., Kvaček et al., 2000, Kvaček and Rember, 2007, Shi et al., 2011).

In this paper, a total of 21 specimens of fossil foliage collected from the upper Miocene of Zhejiang Province, China, are described morphologically and anatomically as a new species, Fokienia shengxianensis sp. nov. These represent the first fossil with a description of the cuticle of Fokienia. Furthermore, paleoecological implications are briefly discussed.

Section snippets

Fossil material and preparation

The fossil leaf material was collected from the Shengxian Formation in Jiahu Village of Tiantai County, and Huangnitang Village of Ninghai County (29° 09′ N, 121° 14′ E: Fig. 1) in Zhejiang Province, eastern China. The formation is dominated by alternating basalts and sediment layers. Fossiliferous layers are exposed in an opencast diatomite mine, which consists of diatomaceous mudstone, shale and argillaceous siltstone. This sediment was originally named the “Xiananshan Formation” (Li and Guo,

Results

  • Family: Cupressaceae Gray, 1822

  • Genus: Fokienia A. Henry et H.H. Thomas, 1911

  • Species: Fokienia shengxianensis He, Sun et Liu sp. nov.

  • (Plate I, 1–7; Plate II, 1–4; Plate III, 1–4; Plate IV, 1–2; Plate V, 1, 3, 5, 7, 9)

  • Synonym: Fokienia notoensis Matsuo. Li and Guo, pp. 288, pl. 135, Fig. 9–9a, 1982.

  • Diagnosis: Foliage branches spread in flattened sprays. Leaves scale-like, dimorphic, decussate and almost in whorls of four. Facial leaves oblanceolate with obtuse apex. Lateral leaves boat-shaped

Comparisons

The present fossil leaves from the Shengxian Formation are scale-like and occur in whorls of four. In gymnosperms, scale leaves appear in families such as Cupressaceae s.s., Taxodiaceae s.s., and Podocarpaceae (de Laubenfels, 1953). Furthermore, leaves in the Curpressaceae s.s. are essentially whorled (de Laubenfels, 1953). Therefore, there is little doubt that the external foliar morphology alone strongly shows that the present fossils are of cupressaceous affinity.

The Cupressaceae s.s.

Acknowledgments

We are grateful to Wenwen Wen and Dr. Li Xiangchuan for their suggestion; Dr. Liang Xiao, Dr. Zhicheng Lin and PhD students Kequn Dao, Baoxia Du for their help in the field. Thanks are also due to S.X. Guo, Z. Kvaček, S. Manchester and Z.K. Zhou for their kindness to provide some information on Ditaxocladus and J. Donaldson for linguistic improvement. This study was supported by the National Basic Research Program of China (973 Program, no. 2012CB822000 to B. Sun), National Natural Science

References (48)

  • D.J. de Laubenfels

    The external morphology of coniferous leaves

    Phytomorphology

    (1953)
  • A. Farjon

    A Monograph of Cupressaceae and Sciadopitys

    (2005)
  • P.A. Gadek et al.

    Relationships within Cupressaceae sensu lato: a combined morphological and molecular approach

    American Journal of Botany

    (2000)
  • S.X. Guo et al.

    Oligocene Sanhe flora in Longjing County of Jilin, Northeast China

    Acta Palaeontologica Sinica

    (2002)
  • S.X. Guo et al.

    Paleocene megafossil flora from Altai of Xinjiang

    Botanical Bulletin of Academia Sinica

    (1984)
  • K. Huzioka

    The tertiary floras of Korea

    Journal of the Mining College of Akita University, Series A

    (1972)
  • S. Ishida

    The Noroshi flora of Noto peninsula, central Japan

    Memoirs of the Faculty of Science Kyoto University: Series of geology

    (1970)
  • H. Jia et al.

    Microstructures of one species of Quercus from the Neogene in Eastern Zhejiang and its palaeoenvironmental indication

    Earth Science Frontiers

    (2009)
  • V.A. Krassilov et al.

    Plant systematics and differentiation of species over Trans-Beringian land connections including a newly recognized cupressaceous conifer Ditaxocladus Guo & Sun

    Bulletin of Geosciences

    (2009)
  • Z. Kvaček et al.

    Calocedrus robustior (Cupressaceae) and Taxus schornii (Taxaceae): two new conifers from the middle Miocene Latah Formation of northern Idaho

    Paleobios

    (2007)
  • Z. Kvaček et al.

    Cones, seeds, and foliage of Tetraclinis salicornioides (Cupressaceae) from the Oligocene and Miocene of Western North America: a geographic extension of the European Tertiary species

    International Journal of Plant Sciences

    (2000)
  • H.L. Li

    Present distribution and habitats of conifers and taxads

    Evolution

    (1953)
  • H.M. Li

    Neogene floras from eastern Zhejiang, China

  • H.M. Li et al.

    Angiospermae

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