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Equisetum Cf. Pratense (Equisetaceae) from the Miocene of Yunnan in Southwestern China and Its Paleoecological Implications Author(S): Yu‐Ling Zhang, David K

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Equisetum Cf. Pratense (Equisetaceae) from the Miocene of Yunnan in Southwestern China and Its Paleoecological Implications Author(S): Yu‐Ling Zhang, David K Equisetum cf. pratense (Equisetaceae) from the Miocene of Yunnan in Southwestern China and Its Paleoecological Implications Author(s): Yu‐Ling Zhang, David K. Ferguson, Albert G. Ablaev, Yu‐Fei Wang, Cheng‐Sen Li, and Lei Xie Source: International Journal of Plant Sciences, Vol. 168, No. 3 (March/April 2007), pp. 351- 359 Published by: The University of Chicago Press Stable URL: http://www.jstor.org/stable/10.1086/510411 . Accessed: 02/04/2015 03:52 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp . JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. The University of Chicago Press is collaborating with JSTOR to digitize, preserve and extend access to International Journal of Plant Sciences. http://www.jstor.org This content downloaded from 159.226.100.224 on Thu, 2 Apr 2015 03:52:13 AM All use subject to JSTOR Terms and Conditions Int. J. Plant Sci. 168(3):351–359. 2007. Ó 2007 by The University of Chicago. All rights reserved. 1058-5893/2007/16803-0009$15.00 EQUISETUM CF. PRATENSE (EQUISETACEAE) FROM THE MIOCENE OF YUNNAN IN SOUTHWESTERN CHINA AND ITS PALEOECOLOGICAL IMPLICATIONS Yu-Ling Zhang,*,y David K. Ferguson,z Albert G. Ablaev,§ Yu-Fei Wang,1,* Cheng-Sen Li,* and Lei Xie* *State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, 20 Nan Xin Cun, Xiangshan, Beijing 100093, People’s Republic of China; yGraduate University, Chinese Academy of Sciences, Beijing 100039, People’s Republic of China; zDepartment of Paleontology, University of Vienna, Althanstrasse 14, A-1090 Vienna, Austria; and §Pacific Oceanological Institute, Russian Academy of Sciences, Vladivostok 600041, Russia Rhizomes with tubers of Equisetum cf. pratense Ehrhart are described for the first time from the late Miocene of Lu¨ he, Yunnan Province, southwestern China. The rhizomes consist of distinct nodes and internodes. The nodes bear one to four bunches of tubers (two to four tubers per bunch). The tubers are elliptical, rounded, ovate, or spindle shaped, and the most distal is mucronate. The occurrence of E. cf. pratense would indicate a wet habitat, and the presence of rhizomes with tubers implies that burial occurred during the late fall or winter, assuming that the fossil and its nearest living relative have similar climatic tolerances and phenology. These finds, combined with the presence of thermophilous genera Cyclobalanopsis, Castanopsis, and Liquidambar and hygrophilous elements Alnus and Taxodioxylon, indicate a warm and humid subtropical climate at Lu¨ he in the Miocene. Keywords: China, Equisetum, Miocene, paleoecology, paleophenology, tubers. Introduction (2004) suggested that E. bogotense is basal within the genus, and all other members of subgenus Equisetum and all species The genus Equisetum L. (horsetails), the sole living repre- of subgenus Hippochaete group into two major sister clades sentative of the class Sphenopsida, contains 15 species with based on the chloroplast rps4 gene and adjacent noncoding herbaceous habit. It has a nearly worldwide distribution ex- sequences. cept for Australia and New Zealand and is commonly associ- Equisetum-like remains preserved as stems, leaves, cones, ated with wet places (Hauke 1993; Kenrick and Crane 1997; rhizomes, and tubers are common in the sediments of the Nor- Page 1997). Equisetum has been considered to be the sister thern Hemisphere from the Carboniferous to the Pleistocene group to all ferns (Kenrick and Crane 1997), the sister group (Brown 1975; McIver and Basinger 1989). Equisetum bryanii to leptosporangiate ferns (Duff and Nickrent 1999), or the Gould with well-preserved stems and leaves, reported from sister group to Marattiopsida (Pryer et al. 2001). the Jurassic of southeastern Queensland, is regarded as the The genus Equisetum is divided into two subgenera: Equi- earliest member of subgenus Equisetum (Gould 1968), while setum L. and Hippochaete Milde (Hauke 1963, 1974, 1978, Equisetum clarnoi Brown, with silicified stem fragments and 1983, 1993; Page 1972; Gifford and Foster 1989; Zhang numerous small roots, found in the Eocene of Oregon, repre- 2004). Subgenus Equisetum comprises eight species that are sents the first conclusive report of subgenus Hippochaete annual, with superficial stomata, regularly branched stems, (Brown 1975). and obtuse cones. Subgenus Hippochaete consists of seven The tuberous rhizomes of Equisetum have been found species that are usually perennial, with sunken stomata, un- in North America in the Cenomanian (upper Cretaceous) branched stems, and apiculate cones (Hauke 1963, 1974, Dakota Formation of Kansas (Skog and Dilcher 1994) and 1978, 1983, 1993). Rhizomes with tubers are common to most the St. Mary River Formation and Edmonton Formation of species of subgenus Equisetum but are never found in subgenus Alberta (Bell 1949), in the Paleocene Ravenscrag Formation Hippochaete (Thome´ 1885; Campbell 1928; Mehra and Bir of Saskatchewan (McIver and Basinger 1989) and the Smoky 1959; Hauke 1963, 1974, 1978; Bir 1978; Watson and Batten Tower Locality (Christophel 1976), and in the Eocene (Lamotte 1990; Doll 2001). 1952) of Barrell’s Springs and Carbon Station of Wyoming Recent molecular studies support the treatment of two (Lesquereux 1878), as well as in the Oligocene to Miocene subgenera within the genus Equisetum in general (Marais Beaverhead Basins in southwestern Montana (Becker 1969). et al. 2003; Guillon 2004). Marais et al. (2003) identified In Europe they have been discovered in early Cretaceous two monophyletic clades corresponding to the two subgenera (Berriasian) Wealden assemblages from England and Germany (except Equisetum bogotense Kunth) based on a combined (Watson and Batten 1990) and Miocene floras of Iceland (Denk analysis of two chloroplast markers, rbcL and trnL-F.Guillon et al. 2005). In Asia they are known from the Jurassic (Bajocian to Bathonian) Utano Formation of Ochi Shimizu-mura and 1 Author for correspondence; e-mail [email protected]. Shimonoseki, Japan (Kon’no 1962); the early Cretaceous of Manuscript received April 2006; revised manuscript received September 2006. China in the Yixian Formation, Beipiao, Liaoning (Wu 1999; 351 This content downloaded from 159.226.100.224 on Thu, 2 Apr 2015 03:52:13 AM All use subject to JSTOR Terms and Conditions 352 INTERNATIONAL JOURNAL OF PLANT SCIENCES Fig. 1 Map of China indicating the Lu¨ he locality of Yunnan. Sun et al. 2001, pp. 70–71; Zhou et al. 2003), the Muling The fossil tubers (33 specimens) and rhizomes with tubers (13 Formation of Jixi Basin, Heilongjiang (Yang 2003), Fuxin For- specimens) of Equisetum are preserved as impressions. They are mation in Liaoning (Chen et al. 1988, pp. 31–32), Huolinghe housed in the National Museum of Plant History of China, Formation, Damoguaihe Formation, and Yinmin Formation Institute of Botany, Chinese Academy of Sciences, Xiangshan, in Inner Mongolia (Deng 1995, pp. 11–14; Deng et al. 1997, Beijing. The morphology was revealed using the degagement pp. 18–20); the middle Eocene (Ablaev et al. 2003) Hunchun technique and was observed under a stereomicroscope. Formation, Jilin in China (Guo 2000); and the Miocene Lawula The herbarium sheets that we examined originated from Formation of Markam and Wulong Formation of Nanmulin PE, Institute of Botany, Chinese Academy of Sciences; For- in Tibet (Xizang) (Geng and Tao 1982; Tao and Du 1987). estry Herbarium of Beijing Forestry University; and field col- In this article, we describe fossil specimens preserved as lections by Yu-Ling Zhang. The habitat and phenology of rhizomes with tubers from a new Miocene locality, Lu¨ he in Equisetum pratense Ehrhart in Songshan Mountain (40°309N, Yunnan, southwestern China, and assign them to Equisetum 115°489E), Beijing, were observed and recorded. cf. pratense Ehrhart. The paleoenvironments of Lu¨he, Yunnan Province, are interpreted using the combined data of sporo- morphs, fossil wood, and other macrofossils. Systematic Description Class—Sphenophytina Material and Methods Order—Equisetales The fossils described here were all collected from the Lu¨ he Family—Equisetaceae lignite mine (25°79–109N, 101°189–229E) of Chuxiong Yi Au- Genus—Equisetum Linnaeus 1753 tonomous Prefecture, Yunnan Province, southwestern China (fig. 1). They were embedded in a gray sandy underclay im- Subgenus—Equisetum 1887 mediately below the lower part of brown coal layer K5, Species—Equisetum cf. pratense Ehrhart 1784 Shigucun beds, Xiaolongtan Formation (BGMRYP 1996; and the 1962 geological report from Geological Team 11 of Description Yunnan Province [GTYN]). The age of the Xiaolongtan For- Rhizomes have distinct nodes (fig. 2A,2H–2J, see arrows a mation is Miocene, based on lithostratigraphic grounds in fig. 2; fig. 3) and internodes (fig. 2I, arrow b). Internodes (BGMRYP 1996). Unfortunately, we lack more precise radio- are 6–10 mm wide and at least 7 cm long, with six to eight metric dating for this. longitudinal ribs on their surface (three to four on each side; Fig. 2 Equisetum cf. pratense. A, Specimen LH1401. B, Specimen LH1511. C, Specimen LH1603. D, Specimen LH1605. E, Specimen LH1405. F, Specimen LH1510. G, Specimen LH1508. H, Specimen LH1411. I, Specimen LH1413. J, Specimen LH1407. a ¼ node; b ¼ internode. Scale bar ¼ 1 cm. This content downloaded from 159.226.100.224 on Thu, 2 Apr 2015 03:52:13 AM All use subject to JSTOR Terms and Conditions This content downloaded from 159.226.100.224 on Thu, 2 Apr 2015 03:52:13 AM All use subject to JSTOR Terms and Conditions 354 INTERNATIONAL JOURNAL OF PLANT SCIENCES Fig. 3 Reconstruction of Equisetum cf. pratense. A, Specimen LH1401. B, Specimen LH1411. C, Specimen LH1407. Scale bar ¼ 1 cm. fig. 2I, arrow b).
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