Studies on the Late Permian Permineralized Tree Fern Psaronius Housuoensis Sp. Nov. from Yunnan Province, Southwest China
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Review of Palaeobotany and Palynology 163 (2011) 247–263 Contents lists available at ScienceDirect Review of Palaeobotany and Palynology journal homepage: www.elsevier.com/locate/revpalbo Research paper Studies on the Late Permian permineralized tree fern Psaronius housuoensis sp. nov. from Yunnan Province, southwest China Ashalata D'Rozario a,⁎, Bin Sun b, Jean Galtier c, Shi-Jun Wang b, Wen-Yi Guo b,d, Yi-Feng Yao b, Cheng-Sen Li b a Department of Botany, Narasinha Dutt College, 129, Bellilious Road, Howrah 711 101, India b State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, Xiangshan, Beijing 100093, PR China c UMR Botanique et Bioinformatique, AMAP, CIRAD, TA A-51/PS2, Boulevard de la Lironde. 34398 Montpellier cedex 05, France d 2 Xiamen Environmental Protection Institute, Xiamen 361006, PR China article info abstract Article history: Psaronius housuoensis sp. nov. is described from the Upper Permian Xuanwei Formation deposits of Housuo Received 2 May 2010 Coal Mine, Fuyuan County, Yunnan Province, southwest China, adding to the marattialean fern community of Received in revised form 1 November 2010 the Cathaysian flora. The permineralized stem shows a polycyclic dictyostele with five leaf traces arranged in a Accepted 2 November 2010 helical manner and exhibiting 2/5 phyllotaxy. Vascular bundles arranged in 6 concentric rings with cauline Available online 10 November 2010 bundles in 5 radial rows. Stem bound externally by a peripheral sclerenchyma sheath. Leaf trace configuration comprised of three vascular bundles resulting from the division of a single originally C-shaped strand. Ground Keywords: Marattiales tissue replaced with coprolites and sediments. Stem surrounded by a thick mantle of roots. Occurrence of Psaronius coprolites within the stem indicates well developed plant–animal interaction during the Late Permian times Late Permian and that the habitat in which Psaronius grew was abounding with diversity of arthropod or the oribatid mite anatomy community preferring the rotting P. housuoensis sp. nov. stem ground tissue as a suitable microenvironment leaf traces for shelter. Evolutionary pattern of leaf trace development is described in the new species and its possible ecological significance. © 2010 Elsevier B.V. All rights reserved. 1. Introduction trunks from Euramerican flora are common (Cotta, 1832; Renault, 1869; Morgan, 1959; Mickle, 1984), or as coal balls (Smoot, 1984; Galtier and The tree fern Psaronius widely distributed during the Late Permian Phillips, 1985; Millay, 1997; Tian et al., 1996; He et al., 2008,) or may (Lopingian) times has greatly attracted the attention of many scientists occur as pyroclastic tuffitic material (Corda, 1845; Rößler and Barthel, because of its unique stelar structures and pattern of leaf trace 1998). Falcon-Lang, 2006 reported the autochthonous occurrence of development. Spanning the time period from earliest Pennsylvanian Psaronius trunks from latest Mid-Pennsylvanian coastal plain deposits in (DiMichele and Phillips, 1977) to Late Permian and -Triassic period Sydney Basin. Globally Psaronius has been recorded from Europe, North (Frentzen, 1914; Hirmer, 1927) they are important elements of the Late America, South America, Southeast Asia and China (He et al., 2008). Carboniferous and Permian flora (Cotta, 1832; Corda, 1845; Zeiller, The late Palaeozoic Cathaysian flora of China has preponderance of 1890; Stenzel, 1906; Morgan, 1959; Phillips et al., 1974, 1985; several marattialean ferns, among which Psaronius is a common Pfefferkorn and Thomson, 1982; DiMichele and Phillips, 1994; Falcon- element. Permineralized trunks of several species of Psaronius com- Lang, 2006; He et al., 2008) of the tropical and subtropical regions of monly occur in the late Permian floral assemblages of China. P. sinensis both the Northern and Southern Hemispheres. These plants had a wide (Sze, 1942, 1947) was reported from the Emeishan Basalt of Weining geographical distribution as evidenced by their common occurrence in County, Sichuan Province; P. hexagonus (Gu and Zhi, 1974) was recorded the Euramerican, Cathaysian and Gondwana flora (Rößler, 2000). In from Xuanwei Formation of Sichuan Province; P. yunnanensis from addition they also grew in varied ecological habitats; some species in the Xuanwei Formation of Kunming City, eastern Yunnan Province (Yang, clastic hygrophile lowlands (Phillips et al., 1985; DiMichele and Phillips, 1986); P. jiangsuensis and P. octogonus from the Longtan Formation of 1994), some in peat forming hygrophile mires while others in the Jiangsu Province (Yao et al., 1994); P. wangii (Tian et al., 1992; Li and Cui, mesophilic soil with periodic drier conditions (Barthel and Weiss, 1997; 1995; Tian and Wang, 1995; Tian et al., 1996), P. cf magnificus (Li and Cui, Falcon-Lang, 2006). Records of Psaronius as permineralized, silicified 1995; Tian et al., 1996)andP. panxianensis (He et al., 2008)from Xuanwei Formation of Panxian County, Guizhou Province. Several isolated pieces of Psaronius sp. root mantles were found in the ⁎ Corresponding author. Department of Botany, Narasinha Dutt College, 129, Bellilious Road, Howrah 711 101, India. volcaniclastic tuffs from Xuanwei Formation of Panxian County, E-mail address: [email protected] (A. D'Rozario). Guizhou Province, China (Hilton et al., 2004). 0034-6667/$ – see front matter © 2010 Elsevier B.V. All rights reserved. doi:10.1016/j.revpalbo.2010.11.002 248 A. D'Rozario et al. / Review of Palaeobotany and Palynology 163 (2011) 247–263 Zhong and Smyth (1997) reported the presence of Psaronius on the tannularia cathavsiana, Fascipteris stena, Gigantopteris dictyophylloides, basis of occurrence of ‘barkinite’ in Late Permian marine-influenced coal Gigantonoclea guizhouensis,andRhipidopsis pani were found from the balls of Changguang, Leping and Shuicheng Basins of southern China. fourth part (57.7 m in thickness) of Xuanwei Formation (Bureau of The genus Psaronius was initially described by Cotta (1832) for Geology and Mineral Resources of Yunnan Province, 1990). segments of permineralized trunks, but presently used to refer to the Anatomical study of specimen no. HS-27 is described in the entire plant. In general habit these plants are similar to the present present paper. tree ferns with an upright unbranched stem, bearing a crown of Since the specimens were highly permineralized and etching with spirally arranged pinnate compound leaves. The stem increases in size hydrofluoric or hydrochloric acid did not prove successful, the ‘peel and complexity from base upwards (Morgan, 1959). At the extreme method’ (Galtier and Phillips, 1999) of examining the specimen was base there are mainly roots and the stem is represented by a small not possible, and so the ‘thin section method’ (Hass and Rowe, 1999) siphonostele, which gradually expands upwards to a polycyclic was employed. The permineralized specimen was cut into several dictyostele at the apex, thus forming an inverted cone shaped thin transverse sections with the help of the rock cutting machine construction. The root mantle consists of an inner zone with roots (SPOJ-300), ground and polished with 100 and 300 grade carborun- of small diameter and an outer zone of larger roots. dum powder on single disc polishing machine and mounted on to The present paper describes the anatomical features of a new glass slides using epoxy resin and triethanolamine. It was further species of Psaronius, P. housuoensis sp. nov. from the Upper Permian, polished with 600 and 1000 grade carborundum powder on a glass Xuanwei Formation deposits of Housuo Coal Mine, Fuyuan County, plate for observation under transmitted and reflected light. Yunnan Province, southwest China, the evolutionary pattern found To determine the phyllotaxy and developmental stages of the leaf therein and possible ecological significance. traces serial sections of the 23-cm length of the stem has been cut and slides prepared. Sixteen such thin section slides were prepared and 2. Materials and methods numbered as HS-27-01 to 16 from top to base. HS-27-01 represents the topmost section and HS-27-16 the basal part. As the material is The materials of Psaronius for the present investigation were collected highly petrified so in the process of preparation of the thin sections as permineralized fossil trunks from Upper Permian deposits of Xuanwei there is every possibility that some layers will be worn out during FormationofHousuocoalmine,FuyuanCounty,YunnanProvince, polishing. Taking into account the limitations of the conditions of southwest China (Fig. 1). About 30 trunk specimens numbering from preservation, study of these sections has been undertaken to reveal HS-01 to HS-30 were collected from the coal seam and cut (Fig. 2). None of the pattern of leaf arrangement and leaf trace development. The slides the specimens cut was found to be similar to the studied specimen HS-27. were observed and studied by Leica DMRE microscope, Leica DM 2500 No other plant taxa were found in association with the tree fern trunks microscope and Orient SMI Stereo microscope and photographed by collected from the coal seam. But fossil plants such as Leidodendron Canon EOS 20D digital camera, Nikon Coolpix 4500 and Leica DFC 420 lepidophloides, Paracalamites stenocostatus, Compsopteris contracta, Loba- camera. Images were adjusted in Adobe Photoshop (V. 7) and plates Fig. 1. Map of Fuyuan County, Yunnan Province, southwest China showing position of Housuo coal mine and locality of fossil