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Vessel-Bearing Stems of Vasovinea Tianii Gen. Et Sp

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Vessel-Bearing Stems of Vasovinea Tianii Gen. Et Sp American Journal of Botany 86(11): 1563±1575. 1999. VESSEL-BEARING STEMS OF VASOVINEA TIANII GEN. ET SP. NOV.(GIGANTOPTERIDALES) FROM THE UPPER PERMIAN OF GUIZHOU PROVINCE,CHINA1 HONGQI LI AND DAVID WINSHIP TAYLOR2 Department of Biology, Indiana University Southeast, New Albany, Indiana 47150 Permineralized gigantopterid stems of Vasovinea tianii Li et Taylor gen. et sp. nov. were collected from the Upper Permian of Guizhou Province, China. They are slender and bear prickles, trichomes, and compound hooks. Internally, the stems have a sparganum cortex, eustele, and secondary xylem. The mesarch protoxylem tracheids have annular to helical thickenings, and metaxylem tracheary elements have scalariform and/or transversely elongated, bordered pits, while those of the sec- ondary xylem have scalariform to circular bordered pits. Importantly, the inner part of the secondary xylem has large vessel elements with foraminate-like perforation plates. The hooks and other morphological and anatomical characteristics are similar to those found in gigantopterids, suggesting that Vasovinea is a member of the Gigantopteridales. The vegetative plant is reconstructed from permineralized stems and Gigantopteris-type leaves based on the anatomical similarities and intimate association. The eustele, secondary xylem, and other features support the placement of the order among the seed plants. Ecologically, Vasovinea is suggested to have been a vine or liana that used compound hooks to climb among the trees in a Permian tropical rain forest. The occurrence of vessels could have been an ef®cient adaptation to allow the slender stems to conduct suf®cient water to the large Gigantopteris-type leaves. Key words: gigantopterids; Gigantopteris; hooks; lianas; paleoecology; Permian; stems; Vasovinea; vessels. Fossil specimens of gigantopterids have been found the distinctive leaves, anatomically preserved reproduc- extensively from the Lower Permian and possibly into tive organs have not been found so that the systematics the lowest Lower Triassic sediments throughout south- of the group has been uncertain. eastern Asia (mostly from China), as well as from several Recently, some permineralized gigantopterid leaves Lower Permian sites in Texas and Oklahoma, USA. This and axes have been reported with many anatomical char- group was ®rst reported by Schenk (1883) from Hunan acteristics that can help in determining the relationships Province, China, and currently includes more than 40 of gigantopterids to other vascular plants. The perminer- taxa (H. Li et al., 1994). The distinctive characteristics alized leaves of the gigantopterid Delnortea abbottiae, of gigantopterids are their large compound or simple reported from Texas, USA, are suggested as being struc- leaves, which are variable in morphology, ranging from turally similar to gnetophytes (Mamay et al., 1988). In oblong to roughly round shape, entire to toothed margin, contrast, the permineralized leaves of Gigantonoclea and simple to complex reticulate venation. Convention- guizhouensis, from the Upper Permian in Guizhou, Chi- ally the group has been classi®ed as ferns (Schenk, 1883) na, exhibit many features similar to those of angiosperms or seed ferns (White, 1912; Asama, 1959; X. Li and Yao, (H. Li and Tian, 1990; H. Li et al., 1994). Two types of 1983). In terms of relationships to other plants, Asama permineralized gigantopterid stems also are found asso- (1974, 1982, 1988) proposed that the simple-leafed gi- ciated with the Guizhou gigantopterid leaves (H. Li, Tay- gantopterids could have given rise to the angiosperms, lor, and Taylor, 1992, 1993). One type is the prickly Acu- while X. Li and Yao (1983) interpreted their reconstruc- leovinea yunguiensis (H. Li and Taylor, 1998), which is tion of the reproductive organs of Gigantonoclea fuki- considered a seed plant. Another type containing vessels enensis as being ``parti-angiosperms'' in nature. Despite has been reported preliminarily (H. Li, Taylor, and Taylor, 1996). 1 Manuscript received 16 July 1998; revision accepted 25 March Before the report by H. Li, Taylor, and Taylor (1996), 1999. the fossil record of vessels could be traced back to the This paper represents part of Hongqi Li's dissertation submitted in partial ful®llment of the requirements for the Ph.D. degree at the Ohio Early Cretaceous. In extant plants, vessels consist of ver- State University, Columbus, Ohio. Hongqi Li thanks Drs. Edith L. Tay- tically linked tracheary elements with perforate end walls lor and Thomas N. Taylor for professional help, and Dr. William A. so that they conduct water much more ef®ciently than Jensen for guidance and encouragement. The authors thank Dr. Cheng- tracheids with imperforate end walls. Vessels are com- sen Li and the Botanical Institute of Beijing for support in fossil-col- monly found in angiosperms and gnetophytes, with rarer lecting; Dr. Paul Kenrick and the staff of the Swedish Museum of Nat- ural History for loan of gigantopterid specimens; and Drs. Sherwin occurrences in non-seed plants (see Discussion). Vessels Carlquist, Kathleen B. Pigg, Leo J. Hickey, and an anonymous reviewer in living seed plants have been classi®ed in two types, for their valuable comments on the manuscript. This research is partially i.e., foraminate vessels in gnetophytes and scalariform supported by the Geological Society of America Research Grant ones (and/or their derived simple form) in angiosperms 5195-93, the Graduate Student Alumni Research Award, Graduate (Bailey, 1944; Carlquist, 1992, 1994, 1996a). Vasovinea School of the Ohio State University, and grants from the Research and University Graduate School of Indiana University, and the Academic tianii is different from the above two types, possessing Affairs of Indiana University Southeast. unique, foraminate-like vessels in the secondary xylem 2 Author for correspondence (e-mail: [email protected]). and a possible scalariform-reticulate vessel in the meta- 1563 1564 AMERICAN JOURNAL OF BOTANY [Vol. 86 xylem. Therefore, the discovery of vessels in gigantop- cm in diameter, bearing compound hooks, prickles, and terids is important not only as the earliest fossil record glandular and tendril-like trichomes. Cortex with paired of vessels, but also in analyzing the systematic relation- vascular traces and sparganum structure in the outer part. ships and the ecological aspects of the group. Eustele with parenchymatous pith and mesarch protoxy- To complement the preliminary study (H. Li, Taylor, lem, consisting of tracheids with annular, helical, or he- and Taylor, 1996), we now provide a comprehensive de- lical-scalariform thickenings. Centripetal metaxylem tra- scription of the permineralized, vessel-bearing gigantop- cheids commonly consisting of one to two layers, while terid stems and establish them as a new taxon, Vasovinea centrifugal metaxylem usually consisting of two to three tianii Li et Taylor gen. et sp. nov. With several lines of layers of tracheary elements; metaxylem tracheary ele- evidence from both permineralized and compression ments with scalariform and/or transversely elongated bor- specimens, we also demonstrate that the new taxon is a dered pits on their lateral walls. Outer portion of the sec- member of the gigantopterids, reconstruct it together with ondary xylem consisting of radial ®les of tracheids and Gigantopteris-type leaves, analyze its possible habit, and smaller vessels. Lateral walls of tracheary elements in brie¯y discuss its systematic relationships to seed plants. secondary xylem exhibiting multiseriate, alternate, and transversely elongate to circular bordered pits. The inner MATERIALS AND METHODS portion of the secondary xylem consisting of large ves- sels with diameter increasing from ;150 to 250 (up to Limestone and mudstone samples with gigantopterid stems and 500) mm towards the primary xylem; vessel elements ver- leaves were collected from a talus pile mined from the upper and middle tically connected by foraminate-like perforation plates on parts of the Xuanwei Formation of the Upper Permian at the Shan-Jian- the long, inclined (usually) to short, almost horizontal Shu site, Yueliangtian Coal Mine in Panxian County, Guizhou, China, (occasionally) end walls, each plate with multiseriate, al- in 1993. Additional information about the locality and stratigraphy can ternate, obliquely elliptical to circular pores without bor- be found in H. Li et al. (1994) and H. Li and Taylor (1998). ders. Homocellular, heteroseriate rays occurring every Permineralized specimens from ®ve limestone samples (L9407, PLY02, PLY03, PLY04, and L9414), and compressed specimens from one to three radial tracheary ®les, consisting of uni- to three mudstone samples (L9426, L9448, and L9449) were used. All bi-seriate xylic rays and multiseriate medullar rays be- samples contain gigantopterid foliage and stem(s) in each, and some tween xylem segments. have additional compound hooks as well as other structures, although not all of these organs from each of the samples are ®gured in this HolotypeÐSlides L9407-C-B2, L9407-C-B16, and paper. Samples PLY03 and PLY04 are small, but each contains a Va- L9407-D-T2. Figs. 1±4, 6. sovinea tianii stem and a piece of Gigantopteris-type leaf. Sample L9407 has numerous pieces of Gigantopteris-type leaves and several ParatypesÐSlides PLY02-C10-1-1, PLY02-E-1, pieces of V. tianii stems. PLY02 is a large sample, weighing more than PLY03±01, PLY03±06, PLY03±07, PLY03±11, PLY03± 8 kg, and contains hundreds of pieces of permineralized (usually gi- 34, and PLY04-B;
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