Research Paper

Stem cambial variants of the Clematis species (Ranunculaceae) in Taiwan

Sheng-Zehn Yang, Po-Hao Chen, Chien-Fan Chen

Published on: 11 November 2021

Page: 526 - 540

DOI: 10.6165/tai.2021.66.526


Studies on cambial variants of Taiwanese species from the genus Clematis (Ranunculaceae) are scarce. This study aimed to investigate and compare the patterns of secondary growth in stems of 22 Clematis taxa, including 19 species and three varieties. The experimental results showed that approximately 27% of the Clematis species in Taiwan had 12 vascular bundles in the stem. In three cambial variants of the genus, the stem was characterized by an irregular conformation, axial vascular elements in segments, and a wedge-like phloem. The rhytidome is composed of periderm and non-conducting phloem and forms either cogwheel-like or continuous segment bark. Eight Clematis species formed arc-like bark, which developed the cogwheel-like rhytidome. Most Clematis species had an indentation in the region of the wider rays, and the wedge-like indentations developed in the interfascicular region of C. henryi var. morii and C. pseudootophora. Narrow rays that occurred in the secondary xylem increased with increasing stem diameter. The vessels dispersed throughout the stem were semi-ring-porous in most species but ring-porous in others. No species had diffuse porous vessels. The vessel restriction pattern was only found in the two species, C. psilandra and C. tsugetorum. The primary xylem ring was located around the pith of C. uncinata var. uncinata, making its pith cavity hexagonal in shape. Five species had the pith cavity feature. This study provides important basic information about Clematis lianas and addresses their taxonomy, which will help to understand their morphology and to ensure the conservation of their diversity.

Keyword: Ranunculaceae, ray indentation, ring bark, scale bark, vessel restriction, wedge phloem

Literature Cited

Angyalossy, V., G. Angeles, M.R. Pace, A.C. Lima, C.L. Dias-Leme, L.G. Lohmann and C. Madero-Vega. 2012. An overview of the anatomy, development and evolution of the vascular system of lianas. Plant Ecol Divers 5(2): 167–182.
DOI: 10.1080/17550874.2011.615574View Article Google Scholar

Angyalossy, V., M.R. Pace and A.C. Lima. 2015. Liana Anatomy: A broad perspective on structural evolution of the vascular system. In: Schnitzer, S.A., F. Bongers, R.J. Burnham and F.E. Putz (eds.) Ecology of Lianas. pp. 253–287. Wiley Blackwell, West Sussex, U.K.
DOI: 10.1002/9781118392409.ch19View Article Google Scholar

Beck, C.B. 2011. An Introduction to Plant Structure and Development: Plant Anatomy for the Twenty-First Century. 2nd edn. Cambridge University Press, Cambridge.

Carlquist, S. 1992. Woody anatomy of selected Cucurbitaceae and its relationship to habit and systematics. Nordic J Bot 12(3): 347–355.
DOI: 10.1111/j.1756-1051.1992.tb01312.xView Article Google Scholar

Carlquist, S. 1993. Wood and bark anatomy of Aristolochiaceae: systematic and habitual correlations. IAWA J 14(4): 341–357.
DOI: 10.1163/22941932-90000588View Article Google Scholar

Carlquist, S. 1995. Wood and bark anatomy of Ranunculaceae (including Hydrastis) and Glaucidiaceae. Aliso 14(2): 65–84.
DOI: 10.5642/aliso.19951402.02View Article Google Scholar

Carlquist, S. 2001. Cambial Variants (Anomalous Secondary Growth). In: Carlquist, S. (ed.) Comparative Wood Anatomy, 2nd edn. pp 271–295. Systematic, Ecological and Evolutionary Aspects of Dicotyledon Wood. Spinger-Verlag Press, Berlin, Germany.

Esau, K. 1958. Plants Anatomy. John Wiley & Sons, Inc. New York, USA.
DOI: 10.1021/ac60137a759View Article Google Scholar

Evert, R.F. 2006. Esau’s Plant Anatomy. Meristems, Cells and Tissues of the Plant Body. Their Structure, Function and Development. Wiley and Sons. Hoboken, NJ, USA.

Ferreira, T. and W. Rasband. 2011. The ImageJ User Guide Version 1.44.

Gregory, M. 1994. Bibliography of Systematic Wood Anatomy of Dicotyledons. IAWA J Suppl 1.

IAWA Committee on Nomenclature. 1964. Multilingual Glossary of Terms Used in Wood Anatomy. Verlagsbuchanstalt Konkordia, Winterthur, Switzerland. p. 186.

Isnard, S., N. Rowe and T. Speck. 2003a. Growth habit and mechanical architecture of the sand dune?adapted climber Clematis flammula L. var. maritima L. Ann. Bot. 91(4): 407–417.
DOI: 10.1093/aob/mcg044View Article Google Scholar

Isnard, S., T. Speck and N.P. Rowe. 2003b. Mechanical architecture and development in Clematis: implications for canalised evolution of growth forms. New Phytol 158(3): 543–559.
DOI: 10.1046/j.1469-8137.2003.00771.xView Article Google Scholar

Mauseth, J.D. 1988. Plant Anatomy. The Benjamin/Cummings Publishing Company, Menlo Park, CA.

Pace, M.R., L.G. Lohmann and V. Angyalossy. 2011. Evolution of disparity between the regular and variant phloem in Bignonieae (Bignoniaceae). Am. J. Bot. 98(4): 602–618
DOI: 10.3732/ajb.1000269View Article Google Scholar

Putz, F.E. and H.A. Mooney. 1991. The Biology of Vines. pp 53–71. Cambridge University Press. UK.

Schweingruber, F.H., A. B?rner and E.D. Schulze. 2011. Ranunculaceae. In: Schweingruber, F.H. et al. (eds.) Atlas of Stem Anatomy in Herbs, Shrubs and Trees. 1: 352–371. Springer-Verlag, Berlin Heidelberg.
DOI: 10.1007/978-3-642-11638-4_2View Article Google Scholar

Sieber, M. and L.J. Kucera. 1980. On the stem anatomy of Clematis vitalba L. IAWA Bulletin 1(1-2): 49–54.
DOI: 10.1163/22941932-90000804View Article Google Scholar

Smith, P. 1928. A comparative study of the stem structure of the genus Clematis with special reference to anatomical changes induced by vegetative propagation. Earth Environ. Sci. Trans. R. Soc. Edinb. 55(3): 643–664.
DOI: 10.1017/S008045680001334XView Article Google Scholar

Yang, S.Z. and P.H. Chen. 2015. Classifying Taiwan lianas with radiating plates of xylem. Taiwania 60(4): 151?159
DOI: 10.6165/tai.2015.60.151View Article Google Scholar

Yang, S.Z. and P.H. Chen. 2016. Cambial variants in the Family Menispermaceae in Taiwan. Am. J. Plant Sci. 7(6): 841–854
DOI: 10.4236/ajps.2016.76080View Article Google Scholar

Yang, T.Y.A. and T.C. Huang. 1996. Ranunculaceae. In: Huang, T.-C. et al. (eds.), Flora of Taiwan, 2nd ed. 2: 694–714. Editorial Committee, Dept. Bot., NTU, Taipei, Taiwan.

Wang, W. and B. Bartholomew. 2007. Clematis. In: Wu, Z.Y., P.H. Raven and D.Y. Hong (eds.) Flora of China 6: 333–360. Science Press & Missouri Botanical Garden.