A. L. Takhtajan

A. S. Peirce

C. J. Quinn

C. Luerssen

C. N. Miller

D. A. Johansen

D. C. Christophel

D. Vogellehner

I. N. Sveshnikova

J. M. Anderson

K. A. Jurasky

K. I. Saiki

K. P. Sporne

M. N. Bose

N. V. Gordenko

Ning Tian

P. Menzel

Q. Ji

R. A. Price

R. A. Stockey

R. Florin

R. Pilger

S. B. Manum

S. E. Schlarbaum

S. L. Zheng

ShaoLin Zheng

T. Ohsawa

V. M. Dörken

W. Dallimore

W. Zhang

Wu Zhang

X. M. Jiang

X. X. Hu

Y. D. Wang

Y. Zhou

YongDong Wang

ZiKun Jiang

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    © The Author(s) 2011. This article is published with open access at Springerlink.com csb.scichina.com   www.springer.com/scp                       *Corresponding author (email: ydwang@nigpas.ac.cn) Letter SPECIAL TOPICS:  Geology February 2012  Vol.57  No.6: 569572  doi: 10.1007/s11434-011-4850-z Occurrence of Sciadopitys-like fossil wood (Coniferales) in the  Jurassic of western Liaoning and its evolutionary implications JIANG ZiKun1,3, WANG YongDong1*, ZHENG ShaoLin2,4, ZHANG Wu2 & TIAN Ning4 1 Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008, China; 2 Shenyang Center of China Geological Survey, Shenyang Institute of Geology and Mineral Resources, Shenyang 110034, China; 3 Graduate University of Chinese Academy of Sciences, Beijing 100049, China; 4 Institute of Palaeontology, Shenyang Normal University, Shenyang 110034, China  Received July 5, 2011; accepted October 8, 2011; published online November 19, 2011  The genus Sciadopitys, containing a single species (S. verticillata), is the only extant representative of the Family Sciadopityaceae (Coniferales), and is a remarkable living fossil. Although some leafy fossils have been ascribed to Sciadopityaceae, fossil xylem material with a close affinity to this family is very rare, and there have been no fossils found showing both pith and primary xylem structures, which are of great importance for wood identification. Thus, it has been difficult to use fossils for the understanding of wood anatomy evolution in the sciadopityaceous plants over geological time. In this note we briefly report on Sciadopitys-like fossil wood found in the Middle Jurassic of western Liaoning, which bears well-preserved Protosciadopityoxylon-type secondary xylem, endarch primary xylem and heterogeneous pith. This is the first report of fossil specimens of the Sciadopityaceae with such detailed preservation of wood structures. The discovery provides precise anatomical evidence for reconstructing the evolu-tionary history and geographical distribution of Sciadopityaceae, as well as contributing to understanding of the fossil diversity of the Jurassic Yanliao Flora in northern China. conifer, Sciadopityaceae, anatomical structure, Tiaojishan Formation, Middle Jurassic, western Liaoning  Citation:  Jiang Z K, Wang Y D, Zheng S L, et al. Occurrence of Sciadopitys-like fossil wood (Coniferales) in the Jurassic of western Liaoning and its evolutionary implications. Chin Sci Bull, 2012, 57: 569572, doi: 10.1007/s11434-011-4850-z    The living fossil Sciadopitys is a monotypic genus with a single species, S. verticillata (Thunb.) Sieb. et Zucc. en-demic in Japan. This species was systematically assigned to the Family Taxodiaceae [1–4]. However, some authors in-sist on the establishment of the new Family Sciadopityaceae, on the basis of morphological, ontogenetic, karyological and immunological data [5–10]. Such a proposal has been accepted by many wood anatomists and botanists, and adopted in a number of publications on gymnosperm classi-fication and systematics [11–14]. There is a close relation-ship between Cupressaceae (including former Taxodiaceae) and Sciadopityaceae in morphological aspects. Most molec-ular studies support the placement of Sciadopityaceae basal to a Cupressaceae plus Taxaceae clade [15,16].  Although Sciadopitys is limited in its current distribution, closely related fossils have been reported from Europe and North America, in addition to Japan. Fossil leaves of Sci-adopitys and its allies are known to occur in the Jurassic to Upper Cretaceous of Europe [10–17]. Other fossil leaves referred to Sciadopityaceae are Sciadopityoides from the Late Jurassic and Sciadopitophyllum in the Late Cretaceous to Paleocene [18–21]. The fossil specimens “double needle leaf ” documented by Florin [22,23] from the Tertiary hy-drogenous coal in Germany were believed to be closely allied to or generically identical to Sciadopitys [24]. “Sci-adopitys-like” leaves also have been described in the Late Jurassic to Early Cretaceous in Europe, including European Russia, such as Tritaenia Maegdefrau et Rudolf [25,26]. Well-preserved seed cones of the Sciadopytiaceae have been documented from the Upper Cretaceous in Japan 570 Jiang Z K, et al.   Chin Sci Bull   February (2012) Vol.57 No.6 [7,27].  The anatomical structures of secondary xylem of the liv-ing species, Sciadopitys verticillata have been studied in details [9,14,28,29]. The unique structure that distinguishes Sciadopitys from other conifers is the cross-field dominated by window-like pits and sub-taxodioid pits with extended pit apertures. Such characteristic anatomical structures are rarely documented in fossil xylem. Jurasky first described Sciadopityoxylon from the Tertiary of Germany [30]. The genus is characterized by uniseriate, separated and rounded bordered pits on radial walls, occasionally with biseriate and opposite pits. The cross-field pits are of both win-dow-like and sub-taxodioid types. There are tracheids on the tangential walls, but spiral thickening and parenchyma are not present. The xylem rays are uniseriate, 1–6 cells high. Ray cells are smooth on the horizontal walls. Resin canals are absent [31,32]. Recently, two sciadopityaceous taxa, Sciadopityoxylon and Protosciadopityoxylon, have been recorded in Early Jurassic and Cretaceous deposits in northeast China [31–33] with generally similar secondary xylem. These two genera show the same window-like and sub-taxodioid cross-field pitting as in the living species, but are different in pits on the radial walls. Pits on the radial walls of Protosciadopityoxylon are uniseriate and close, or biseriate and alternate, while Sciadopityoxylon are uniseri-ate and separate, or biseriate and opposite. Pits on the radial walls of living Sciadopitys are uniseriate, seldom crowded [9]. The extant species S. verticillata is characterized by star-shaped pith and endarch primary xylem [9,29]. Pre- viously, these anatomical features have been entirely un-known in fossil material. Here, we report on new fossil co-niferous wood with Sciadopytiaceae affinity from the Juras-sic of western Liaoning, China, characterized by having well preserved pith, primary xylem and secondary xylem. The specimens were found from the Middle Jurassic Tiaojishan Formation in the Changgao area, Beipiao City of western Liaoning. The Tiaojishan Formation is widely dis-tributed in western Liaoning and northern Hebei and is rep-resented by intermediate lava and pyroclastic rock, with intercalated basic volcanic rocks and sedimentary deposits. This formation has three sedimentary interbeds containing well preserved fossil plants. A number of plant fossils have been reported from this formation, including leaf impres-sions and compressions, seeds and fruits, permineralized fern rhizomes and fossil wood [34]. According to plant fos-sil assemblages and isotopic dating [35], the Tiaojishan Formation is considered to be Middle Jurassic in age. Three well-preserved specimens in our collection belong to the new fossil wood taxon (PB21412–PB21414). The wood bears pith, primary xylem and secondary xylem. The pith is heterogeneous and irregularly stellate in shape (Fig-ure 1a), which consists of parenchyma cells and small number of scattered sclerenchyma cells (Figure 1b). The most remarkable feature of the pith is the presence of dark nodules among the parenchyma cells (Figure 1c, arrow). These nodules are composed of clustered sclerenchyma cells. The primary xylem is endarch with metaxylem grow-ing exocentrically. The radial wall pits on secondary xylem are of the transitional type; where uniseriate, the tracheids are circular and closely arranged (Figure 1d, left arrow); where biseriate, the tracheids are alternate and circular or polygonal (Figure 1d, right arrow). The cross-field pits contain both sub-taxodioid pits with extended pit aperture, and window-like pits. (Figure 1e,f). The ray cells are nor-mally uniseriate, in places biseriate, with a height of 2–26 cells, and longitudinally elliptical in shape. According to the anatomical structures of the secondary xylem, namely the transitional type pitting of the radial wall and mixed window-like pits and sub-taxodioid cross-field pits with extended pit apertures [14], the secondary xylem of our new fossil wood type should belong to the Protosci-adopityoxylon-type of the Sciadopityaceae (Coniferales). The presence of detailed anatomical structures of pith and primary xylem enables us to establish a new morphogenus for the present specimens (detailed description to be pub-lished separately). It is the first time that Sciadopitys-like fossils with pith, primary xylem and secondary xylem have been found in the world. The present specimen has no doubt close systematic affinities to Sciadopitys, Sciadopityoxylon and Protosciadopityoxylon because it resembles, in many respects, the living genus and is generally similar to all three genera in the secondary wood structure. Two other coniferous genera, Scotoxylon (Seward et Bancroft) Vo-gellehner and Yokoxylon (Holden) Vogellehner [36], may be comparable with the specimens from western Liaoning, in having pith, primary and secondary xylem. The three genera share several characteristics in the primary xylem, but are different in pith and secondary xylem structures, especially in the features of cross-field pits. The secondary xylem of Scotoxylon is of the Protojuniperoxylon-type, whereas that of Yokoxylon is of the Protocedroxylon-type. They are not related to Sciadopityaceae with Sciadopitys- (or Sci-adopityoxylon- and Protosciadopityoxylon-) type secondary xylem. The piths of both Scotoxylon and Yorkoxylon consist of homogeneous parenchyma cells, whereas the pith of the new morphogenus from Liaoning is heterogeneous and con-sists of parenchyma cells and scattered sclerenchyma cells. Comparing the new genus with Sciadopityoxylon and Pro-tosciadopityoxylon, their secondary xylem is almost the same, especially the cross-field pitting, except for some minor differences in radial wall pits with Sciadopityoxylon. Pith and primary xylem structures of the new morphogenus from Liaoning are similar to Sciadopitys, namely that both genera show star-shaped pith and endarch primary xylem, which indicates a more close affinity. However, due to the lack of pith and primary xylem structures in Sciadopityoxy-lon and Protosciadopityoxylon, further comparisons among them are currently impossible. It is likely that some speci-mens referred to the Protosciadopityoxylon-type may be found to belong to the new morphogenus from Liaoning, if   Jiang Z K, et al.   Chin Sci Bull   February (2012) Vol.57 No.6 571  Figure 1  Sciadopitys-type fossils from the Middle Jurassic in Western Liaoning. a, Transverse section showing pith (Pi), primary xylem (Px) and second-ary xylem (Sx), the arrow showing primary xylem. Scale bar = 1 mm. b, Transverse section showing parenchyma cells and small number of scattered scle-renchyma cells, the arrow indicating primary xylem and sclerenchyma cell. Scale bar = 100 μm. c, Radial section with nodules consists of clustered scleren-chyma cells (with arrows), surrounded by radial parenchyma cells. Scale bar = 500 μm. d, Radial section with radial wall pits uniseriate (left arrow), biseriate and alternate (right arrow) in arrangement (transitional type). Scale bar = 50 μm. e, f, Radial section showing cross-field pits (e: sub-taxodioid, scale bar= 62.5 μm; f: window-like, scale bar = 25 μm). Specimens and slides are preserved in Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences. ever similar heterogenous pith and endarch primary xylem are detected.  The new morphogenus from Liaoning is similar to Pro-tosciadopityoxylon in that it has transitional type tracheid pitting in the secondary xylem. Hence, it is relatively primi-tive, whereas the xylem structure of Sciadopityoxylon and Sciadopitys both belong to the living type. From a paleobio- geographic viewpoint, it is interesting that although the ex-tant Sciadopitys is endemic in Japan, its ancestors may have existed in the Early and Middle Jurassic in China, flourished in the Jurassic and Cretaceous, and perhaps one of the groups then migrated from China to Western Europe and northwestern Canada. Because of significant changes in palaeoclimate and palaeoenvironment, sciadoptyaceous conifers became extinct in China, Europe and North Amer-ican, but still survived in Japan since the Cretaceous [10]. The new Sciadopitys-type fossil wood from the Middle Ju-rassic of western Liaoning provides significant anatomical evidence for understanding the evolutionary link of sci-adopityaceous wood fossils from the Early and Middle Ju-rassic, to Tertiary and living species. The discovery also increases our understanding of the fossil diversity of the Jurassic Yanliao Flora in northern China. We thank Prof. Zhou Zhiyan (NIGPAS, Nanjing) for carefully reading the manuscript and providing suggestions for improvements. Dr. Mao Limi (NIGPAS, Nanjing), Dr. KazuoTerada (Fukui Prefectural Dinosaur Mu-seum, Japan) and Dr. Marc Philippe (Université Claude-Bernard Lyon 1, France) are thanked for providing references and Dr. Zhang Yi (Shenyang Normal University) for help with photography. This work was supported by the Knowledge Innovation Program of Chinese Academy of Sciences (KZCX-2-YW-154), the National Basic Research Program of China (2012CB822000 and 2006CB701401), the National Natural Science Foundation of China (40972008, 40472004 and J0930006) and UNESCO- IUGS Project IGCP 506. 1 Pilger R. Die Natürlichen Pflanzenfamilien. 2nd ed. Leipzig: W. Engelmann, 1926. 121–403 572 Jiang Z K, et al.   Chin Sci Bull   February (2012) Vol.57 No.6 2 Dallimore W, Jackson A B, Harrison S G. A Handbook of the Conif-erae and Ginkgoaceae. 4th ed. New York: St. Martin’s Press, 1966. 1–729 3 Sporne K P. The Morphology of Gymnosperms. 2nd ed. London: Hutchinson, 1974. 1–29 4 Miller C N. Origin and Evolution of Gymnosperms. New York: Co-lumbia University Press, 1988. 448–486 5 Schlarbaum S E, Tsuchiya T. Bot Gaz, 1985, 146: 264–267 6 Price R A, Lowenstein J M. Syst Bot, 1989, 14: 141–149 7 Saiki K I. Am J Bot, 1992, 79: 989–995 8 Luerssen C. Grundzüge der Botanik. 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Occurrence of Sciadopitys-like fossil wood (Coniferales) in the Jurassic of western Liaoning and its evolutionary implications

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Occurrence of Sciadopitys-like fossil wood (Coniferales) in the Jurassic of western Liaoning and its evolutionary implications

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