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Anatomy of the Late Devonian Sphenopsid Rotafolia Songziensis

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Anatomy of the Late Devonian Sphenopsid Rotafolia Songziensis Anatomy of the Late Devonian Sphenopsid Rotafolia songziensis , with a Discussion of Stelar Architecture of the Sphenophyllales Author(s): De‐Ming Wang, Shou‐Gang Hao, Qi Wang, and Jin‐Zhuang Xue Source: International Journal of Plant Sciences, Vol. 167, No. 2 (March 2006), pp. 373-383 Published by: The University of Chicago Press Stable URL: http://www.jstor.org/stable/10.1086/499115 . Accessed: 02/04/2015 03:18 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 03:18:06 AM All use subject to JSTOR Terms and Conditions Int. J. Plant Sci. 167(2):373–383. 2006. Ó 2006 by The University of Chicago. All rights reserved. 1058-5893/2006/16702-0020$15.00 ANATOMY OF THE LATE DEVONIAN SPHENOPSID ROTAFOLIA SONGZIENSIS, WITH A DISCUSSION OF STELAR ARCHITECTURE OF THE SPHENOPHYLLALES De-Ming Wang,*,y Shou-Gang Hao,1,* Qi Wang,z and Jin-Zhuang Xue* *Key Laboratory of Orogenic Belts and Crustal Evolution, Department of Geology, Peking University, Beijing 100871, China; yInstitute for Earth Sciences, University of Graz, A-8010 Graz, Austria; and zKey Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China A previous study of the Late Devonian (Famennian) sphenopsid Rotafolia songziensis Wang, Hao, and Wang provided detailed descriptions of the morphology and a sketchy illustration of a three-ribbed primary xylem. The present anatomical data show that the protostele of this plant also has four-ribbed primary xylem of exarch maturation. Located at the tip of each xylem rib, the tracheids of the protoxylem strand bear helical wall thickenings. Tracheids of metaxylem and secondary xylem possess scalariform pits and/or bordered pits. Ray cells seldom occur in secondary xylem. In contrast to fertile organs demonstrating great diversity in bract shape, structure, and sporangiophore number, the stelar architecture of the Sphenophyllales is consistent in ribbed primary xylem of fundamentally exarch maturation, although secondary xylem indicates moderate structural variations. Stelar architecture of the Sphenophyllales is compared with that of basal euphyllophytes and lycophytes. The position of peripheral protoxylem strands in the Sphenophyllales and Iridopteridales corresponds to the presence of leaves or leaf precursors. The origination of these strands differs from that of radiate protoxylem strands in the Aneurophytales. Keywords: Rotafolia songziensis, Late Devonian, stelar architecture, sphenopsid, Sphenophyllales. Introduction referring to advanced trimerophytes, aneurophytalean pro- gymnosperms, and some early seed plants and the latter Phylogenetic analyses (Kenrick and Crane 1997a, 1997b; referring to some primitive ferns (e.g., iridopteridalean clado- Doyle 1998) recognized two major monophyletic lineages xylopsids) and early sphenopsids. within the fossil and living vascular plants (tracheophytes). The sphenopsids (also known as equisetophytes, horsetails, One lineage is the lycophytes (i.e., Lycophytina), and the articulates, or arthrophytes) represent one distinctive clade of other is the euphyllophytes (i.e., Euphyllophytina), which are land vascular plants that are characterized by whorled ar- two land-plant sister groups. The lycophytes contain leafless rangement of fertile units and highly reduced vegetative zosterophylls and derived microphyllous lycopsids (club- leaves. The Sphenopsida comprise two major orders, i.e., the mosses). They are characterized by exarch primary xylem Sphenophyllales and the Equisetales. The extinct Sphenophyl- with multiple peripheral protoxylem strands around the cy- lales appeared as early as the Late Devonian, reached their lindrical protostele. Among them, the Lycopodiaceae and maximum development in the Late Carboniferous, and then the Drepanophycales (prelycopsids) possess distinctive ribbed disappeared by the Early Permian. The only extant element primary xylem. The euphyllophytes include ancestral trimer- of the sphenopsids is the genus Equisetum, which belongs ophytes and descended ferns, sphenopsids, progymnosperms, to the Equisetales. A novel sphenophyllalean plant Rotafolia and seed plants. The stelar architecture of this lineage is dif- songziensis (Wang et al. 2005) came from the Late Devonian ferent from and more diversified than that of the lycophytes. (Famennian) Xiejingsi Formation, southwestern Hubei Prov- For example, the trimerophytes have protostele with cen- ince, China. Former study concerned detailed morphological trarch primary xylem, primitive ferns (e.g., cladoxylopsids) description of its branches, vegetative leaves, strobili, and possess actinostele or plectostele with mesarch primary xy- a sketchy illustration of a three-ribbed primary xylem. We re- lem, the sphenopsids bear ribbed or highly separated primary cently obtained additional material of this plant from the xylem of exarch or mesarch/endarch development, aneuro- same locality; thus, it is possible to provide more information phytalean progymnosperms contain ribbed primary xylem of of the internal structure in R. songziensis and compare the mesarch maturation, and seed plants are often provided with stelar architecture of the Sphenophyllales with that of the Iri- eustele. According to Beck and Stein (1993), the euphyllo- dopteridales, Aneurophytales, Equisetales, and Lycophytina. phytes could be anatomically divided into ‘‘radiate protoxy- lem’’ and ‘‘permanent protoxylem’’ groups, with the former Material and Methods 1 Author for correspondence; e-mail [email protected]. The Huangkuang section is situated in Liujiachang town, Manuscript received July 2005; revised manuscript received October 2005. Songzi district, southwestern Hubei. The Devonian strata at 373 This content downloaded from 159.226.100.224 on Thu, 2 Apr 2015 03:18:06 AM All use subject to JSTOR Terms and Conditions 374 INTERNATIONAL JOURNAL OF PLANT SCIENCES this section were divided in ascending order into Yuntaiguan ous sporangia; elongate-cuneate bract bearing a distal and (Givetian of the Middle Devonian), Huangjiadeng (Frasnian many lateral elongate segments. Pendulous elongate sporangia of the Late Devonian), and Xiejingsi (Famennian of the Late abaxially attached to base of bract at the same level. Proto- Devonian) formations. Most anatomical materials of Rotafo- stele comprising three- [sometimes four]-ribbed primary xylem lia songziensis were collected from the lower part of the and radial secondary xylem. Primary xylem maturation ex- Xiejingsi Formation, ca. 2 m below the horizon where Sub- arch, with protoxylem strands at tips of xylem ribs. [Ray cells lepidodendron songziense (Wang et al. 2003) was preserved. rare in secondary xylem.] A few specimens were obtained from the same formation, Tizikou section, Maohutang village, Yidu district, south- Species—Rotafolia songziensis (Feng) Wang, western Hubei. A detailed record of the locality of the Hao, and Wang 2005 emend. Huangkuang and Tizikou sections was presented by Feng (1984) and Wang et al. (2003, 2005). Emended specific diagnosis. Same as for generic diagno- At the Huangkuang section, the fusainized compressions sis. Axes up to 20 cm long, with internodes 1.5-(2.2)-4.8 cm are common in a thin bed and are preserved in hard argilla- long, vegetative axes 0.8-(3.0)-8.3 mm in diameter, and ceous sandstones or silty mudstones of slightly pink or black- branching at angles of 45°–85°. Axial spines 1.0-(2.0)-2.8 gray color. At the Tizikou section, the fusainized material mm long and ca. 0.2 mm wide at base. Wedge- or fan-shaped came from soft silty mudstones of black-gray color. Occa- vegetative leaves six per node of axis, 5.0-(14.5)-24.0 mm sionally, the axes were preserved as limonitic permineraliza- long and 2.0-(11.0)-18.0 mm wide as a whole. Each vegeta- tions (fig. 2a). tive leaf equally or unequally dividing at angles of 20°–80°, In order to know the details of xylem strand and tracheids, two to four times. Below first division, leaf bases 2.7-(6.0)- the fusainized axes were removed from the rock matrix and 9.0 mm long and 0.4-(0.8)-1.5 mm wide. Fertile axes below mounted on a stub for examination with a scanning electron strobili anisotomous at 30°–45°, up to 7.8 cm long and 2.7- microscope (SEM; Amary 1910FE) at 5 kV (figs. 4, 5). In (3.8)-6.4 mm in diameter. Strobili 2.9-(4.9)-8.5 cm long and most examples, the extraxylary tissues were not contained in 1.0-(1.6)-2.2 cm wide, with internodes of strobilar axes 0.5- the axes; therefore, only the xylem column was embedded (0.8)-1.6 cm long. Each strobilus bearing up to 16 whorls of in resin, sectioned at less than 1-mm intervals, ground, and fertile units. Fertile units 6.4-(8.8)-11.0 mm long and 2.0–3.2 then polished to a thickness where the sections could be mm wide and inserted at ca. 60° or 90° to strobilar axis. observed directly in transmitted light. Six and 11 xylem Bract 5.0–6.7 mm long and ca. 2.0 mm wide, with distal and columns were cut into 32 transverse (A01–09,B01–08,C01–03, lateral segments 1.5–8.0 mm long and ca. 0.2 mm wide and D01–03,E01–04,H01–05) and 15 longitudinal (I01–03,J01–02,K 2.0-(3.1)-3.8 mm long and 0.2–0.3 mm wide, respectively. 01–02, L, M, N, O, P, Q, R, S) thin sections, respectively, for Ten to 18 lateral segments attached to bract at 65°–85°.
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