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Diversity Variation, Distribution Pattern, and Evolutionary Implicat +Model PALWOR-305; No. of Pages 20 ARTICLE IN PRESS Available online at www.sciencedirect.com ScienceDirect Palaeoworld xxx (2015) xxx–xxx A systematic overview of fossil osmundalean ferns in China: Diversity variation, distribution pattern, and evolutionary implications a,f,g b,c,∗ d b e Ning Tian , Yong-Dong Wang , Man Dong , Li-Qin Li , Zi-Kun Jiang a College of Palaeontology, Shenyang Normal University, Shenyang 110034, China b Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008, China c Key Laboratory of Economic Stratigraphy and Palaeogeography, Chinese Academy of Sciences, Nanjing 210008, China d College of Geosciences, Yangtze University, Wuhan 430100, China e Chinese Academy of Geological Sciences, Beijing 100037, China f State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008, China g Key Laboratory for Evolution of Past Life in Northeast Asia, Ministry of Land and Resources, Shenyang 110034, China Received 7 August 2014; received in revised form 9 December 2014; accepted 12 May 2015 Abstract The order Osmundales is a unique fern taxon with extensive fossil records in geological past. Diverse osmundalean fossils have been reported from China, ranging in age from the Late Palaeozoic to the Cenozoic. Most of them are based on leaf impressions/compressions, but permineralized rhizomes are also well documented. In this study, we provide a systematic overview on fossil osmundalean ferns in China with special references on diversity variations, distribution patterns, and evolutionary implications. Fossil evidence indicates that this fern lineage first appeared in the Late Palaeozoic in China. The Late Triassic to Middle Jurassic interval was the radiation stage. From the Late Jurassic onward, fossil diversity declined rapidly. Cenozoic osmundalean taxa are represented by the relict species of Osmunda. Geographically, osmundalean fossils are found from both the Northern and Southern phytoprovinces of China, though variations are documented for geographical ranges. The Chinese fossil records cover almost all important stages for the macroevolution of the Osmundales, and contribute to further understanding of evolutionary processes of this peculiar fern lineage. © 2015 Published by Elsevier B.V. on behalf of Nanjing Institute of Geology and Palaeontology, CAS. Keywords: Osmundales; Fossil record; Diversity variation; Distribution pattern; Macroevolution; China 1. Introduction 2006; Schuettpelz and Pryer, 2007; Hennequin et al., 2008). The extant Osmundales are represented mainly by four genera Osmundales, a unique group among the oldest existing ferns, within the family Osmundaceae, including Osmunda, Todea, consist of two major lineages, including the extinct family Leptopteris, and Osmundastrum (Metzgar et al., 2008). The Guaireaceae and the family Osmundaceae with both living and extensive fossil records show, however, that this order had a fossil representatives. This fern clade is generally considered much greater distribution and higher diversification in the geo- as an intermediate between eusporangiate and leptosporan- logical past (Banerji, 1992; Ash and Morales, 1993; Tidwell and giate ferns (Tidwell and Ash, 1994). It is proposed to be the Ash, 1994; Kiritchkova et al., 1999; Collinson, 2001; Bodor and stem-group of the Polypodioidae and sister to the remaining lin- Barbacka, 2008; Taylor et al., 2009). Among these records, over eages of leptosporangiate ferns based on phylogenetic analyses 80 species ascribed to 14 genera of fossil osmundalean rhizomes (Hasebe et al., 1995; Pryer et al., 2001, 2004; Schuettpelz et al., have been documented worldwide, ranging from the Permian to the Cenozoic in age (Tian et al., 2008). In China, limited living members of this fern order are distributed, including Osmunda ∗ Corresponding author: Nanjing Institute of Geology and Palaeontology, Chi- and Osmundastrum mainly in southern China (Wu, 1992; Wang nese Academy of Sciences, Nanjing 210008, China. Tel.: +86 25 8328 2221; and Wang, 2001); as a contrast, abundant fossil osmundalean fax: +86 25 8335 7026. taxa have been reported from the Palaeozoic to the Cenozoic E-mail address: [email protected] (Y.D. Wang). http://dx.doi.org/10.1016/j.palwor.2015.05.005 1871-174X/© 2015 Published by Elsevier B.V. on behalf of Nanjing Institute of Geology and Palaeontology, CAS. Please cite this article in press as: Tian, N., et al., A systematic overview of fossil osmundalean ferns in China: Diversity variation, distribution pattern, and evolutionary implications. Palaeoworld (2015), http://dx.doi.org/10.1016/j.palwor.2015.05.005 +Model PALWOR-305; No. of Pages 20 ARTICLE IN PRESS 2 N. Tian et al. / Palaeoworld xxx (2015) xxx–xxx deposits. A couple of such fossil taxa have been investigated Burakova (Wang et al., 2005; Deng, 2007). Some other foliage in some details, e.g., Todites and fossil rhizomes (Wang et al., taxa referable to the Osmundales have never been documented 2005; Tian, 2011). However, the diversification and distribution in China so far, such as Anomopteris Brongn., Phyllopteroides features of this fern group are still scarcely considered. Medwell, and Cacumen Cantrill et Webb (Cantrill and Webb, In this paper, we review the fossil osmundalean ferns in 1987; Banerji, 1992; Cantrill and Nagalingum, 2005; Taylor China on the basis of known fossil records, with an emphasis et al., 2009). on its diversity variations, distribution patterns, and evolutionary Osmundaceous pinnae are generally characterized by a implications. The scope of this review focuses mainly on macro- pinniform venation; however, several specimens with simple fossil with an exclusion of these osmundalean dispersed spores. reticulated veins were previously assigned to the Osmundaceae This represents the first comprehensive analysis and overview (Osmundales) in China, such as Abropteris yongrenensis Li et on the osmundalean fossils in China, and provides new clues Tsao and Reteophlebis simplex Li et Tsao (Li et al., 1976; Wu, for understanding the origin, radiation, and development of this 1982). In fact, Reteophlebis simplex is a synonym of Cynepteris fern lineage. lasiophora Ash, which was first reported from the Upper Tri- assic Chinle Formation of New Mexico, USA (Ye et al., 1986). 2. Materials and methods The latter was ascribed to a well-defined family Cynepteridaceae (Ash, 1970). The systematic affinity of Abropteris yongrenensis Materials for this study involve all available published data remains poorly understood. However, it shows a close similar- concerning fossil Osmundales in China, such as research papers, ity to Reteophlebis simplex (C. lasiophora) in many characters, monographs and atlases of fossil plants. These fossils mainly fall and may represent an otherwise unknown stock of cynepterida- into two types, i.e., compression/impression foliage and per- ceous plants. It is noted that the Cynepteridaceae is proposed to mineralized rhizome. For each type of the data, the systematic be closely related to the fern family Schizaeaceae (Ash, 1970; attribution, geographical origin, and geological source are com- Axsmith, 2009). piled. Fossil locality information is plotted in a geographical map to show the distribution patterns. The stratigraphic data 3.1.1. Genus Todites Seward emend. Harris are arranged into a time framework, e.g., Early Permian (P1), The genus Todites was erected by Seward (1900) based on Late Permian (P2), Early Triassic (T1), Middle Triassic (T2), materials from Yorkshire, and was then emended by Harris etc. on the basis of up-dated biostratigraphic evidence. Selected (1961). It was named for fossil leaves that morphologically sketch drawings and illustrations of some representative fossil resemble the extant Todea. The sporangia of Todites are always taxa referred to the Osmundales are given, based on figures, borne along the veins on the dorsal side of the fertile pinnae, plates, and descriptions from the original publications. which are morphologically similar to the vegetative ones (Harris, 1961; Wang et al., 2005). The annulus of Todites is 3. Fossil record and diversity of Osmundalean ferns in always strongly apical, covering the entire apical region of China the sporangium (Harris, 1961; Hewitson, 1962). Todites first appeared in the Late Permian, with fertile pinnae bearing in Our general analysis indicates that the earliest record of situ spores from the Upper Permian in Russia (Radcenko,ˇ Osmundales in China dates back to the Late Palaeozoic (Gu and 1955; Naugolnykh, 2002). It is the most common genus Zhi, 1974; Li, 1983). During the Mesozoic, the China territory of the Mesozoic osmundaceous plants in China, and about is divided into two phytoprovinces, i.e., the Northern Phyto- 17 species have been reported (Wang et al., 2005), mainly province (NPP) and Southern Phytoprovince (SPP), delimited by represented by T. shensiensis, T. denticulatus, T. goeppertianus, the Kunlun-Qinling-Dabie Mountain Range (Sun et al., 1995a,b; T. williamsonii, T. princeps, and T. scorebyensis (Fig. 1). The Zhou, 1995). Osmundalean ferns flourished with a high diversity earliest fossil record of Todites in China is from the Middle in both phytoprovinces during the Mesozoic. Several Cenozoic Triassic (e.g., Todites shensiensis from the Middle Triassic of fossil taxa of the family are also reported. The following is a Shaanxi and Inner Mongolia) (Sze, 1956; Zhang, 1976; Huang summary for the diverse type of osmundalean
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