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What Is Special About Plectocomia Himalayana Griff. (Calamoideae, Plectocomiinae)?

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What Is Special About Plectocomia Himalayana Griff. (Calamoideae, Plectocomiinae)? Blackwell Publishing LtdOxford, UKBOJBotanical Journal of the Linnean Society0024-4074The Linnean Society of London, 2006? 2006 151? 8391 Original Article MECHANICAL PROPERTIES AND BRANCHING IN PLECTOCOMIA HIMALAYANA S. ISNARD Botanical Journal of the Linnean Society, 2006, 151, 83–91. With 6 figures The Palms Guest edited by William J. Baker and Scott Zona Biomechanics and development of rattans: what is special about Plectocomia himalayana Griff. (Calamoideae, Plectocomiinae)? SANDRINE ISNARD* Botanique and Bioinformatique de l’Architecture des Plantes, UMR5120 CNRS, TA40/PS,2 Boulevard de la Lironde, F-34398 Montpellier cedex 5, France Received April 2005; accepted for publication November 2005 Mechanical and morphological studies of Plectocomia himalayana (subtribe Plectocomiinae) revealed characteristics that differ strongly from species of subtribe Calaminae (Calamus and Daemonorops). In species of Calaminae tested previously, the contribution of the leaf sheath drastically increases stiffness in juvenile axes and towards the apex of older plants. In P. himalayana the relative contribution of the leaf sheath to axis stiffness is less and leaf sheath senescence does not strongly reduce axial stiffness as observed in Calamus and Daemonorops. Natural aerial branch- ing, only described in Korthalsia and Laccosperma among rattans, is common in P. himalayana. Aerial branching and adventitious roots occur frequently along old stems allowing autonomy of stems, following mechanical injury and promoting vegetative propagation. The climbing habit is known to have evolved at least twice within the Calam- oideae. The results observed here suggest that climbing habits may differ in detail and that different ‘climbing strat- egies’ may have evolved within the subfamily Calamoideae resulting from: (1) variable stem flexibility, (2) the variable mechanical role of the leaf sheath (Calamus–Daemonorops) and (3) production of branches and aerial roots conferring a higher degree of architectural plasticity (Plectocomia). © 2006 The Linnean Society of London, Bota- nical Journal of the Linnean Society, 2006, 151, 83–91. ADDITIONAL KEYWORDS: adventitious roots – branching – Calamus – climbing palms – Daemonorops – layering – leaf sheath – mechanical properties. INTRODUCTION (Baker, Dransfield & Hedderson, 2000a; Baker, Hed- derson & Dransfield, 2000b). According to recent phy- Climbing growth forms have appeared several times logenetic studies, the climbing growth form could have during palm evolution and in two of the five currently multiple origins within the Calamoideae, appearing at recognized subfamilies (Dransfield et al., 2005). In the least twice during evolution (Baker et al., 2000a,b). A Arecoideae climbers include Chamaedorea elatior current project on climbing palms aims to characterize Mart. (tribe Chamaedoreeae), Dypsis scandens J. the mechanical properties and development of species Dransf. (tribe Areceae) and seven species of the genus belonging to different subfamilies and subtribes. The Desmoncus (tribe Cocoseae). The Calamoideae objective is to understand: (1) how such relatively includes 560 climbing species in 13 genera and repre- large-bodied plants lacking secondary growth have sents the most diversified subfamily in this respect adopted a scandent habit, (2) whether different origins of the climbing growth form in the family result in dif- ferent ‘climbing strategies’ and (3) if there are struc- *E-mail: [email protected] tural, mechanical characteristics that can explain the © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 151, 83–91 83 84 S. ISNARD evolutionary success of the subfamily Calamoideae and unusual in palms in general. Among rattans, and most particularly the subtribe Calaminae. aerial branching has only been described for Korthal- Recent studies have shown that despite overall simi- sia (Calamoideae, Korthalsiinae), which is known to larities, different climbing palms can show variable branch frequently in the canopy (Dransfield, 1978; mechanical architectures linked to the climbing habit Uhl & Dransfield, 1987). Aerial branching is rare in (Rowe, Isnard & Speck, 2004). This variation is based arborescent palms and Laccosperma although some around the potentially variable mechanical properties species of Hyphaene and Nypa do develop dichoto- of the inner stem and external leaf sheath of the entire mous branching (Tomlinson, Zimmermann & Simp- axis, and how such properties may vary during devel- son, 1970; Dransfield, 1978). Branch formation is opment. Species tested within Calamus and Daemon- particularly interesting in terms of the ability of a orops share a broadly comparable mechanical feature species to establish itself in the canopy, survive with dicot lianas represented by a decrease of struc- mechanical or hydraulic failure, and offset the life- tural Young’s modulus, Estr in old parts of the climbing span limitation resulting from hapaxanthy. This axis. Whereas in many dicot lianas tested (Rowe & study investigates the morphology and mechanical Speck, 1996, 1998; Isnard, Speck & Rowe, 2003a), architecture of P. himalayana and its ability to branch such changes result from highly sophisticated devel- in the canopy and produce adventitious roots. opment of the vascular cambium. In the species of Cal- amus and Daemonorops tested, the drop in E along str MATERIAL AND METHODS the axis was a consequence of the senescence, decay and eventual loss of the outer relatively rigid leaf Observations and measurements were carried out in sheath component of the axis. In two climbing species the Xishuangbanna autonomous prefecture, Yunnan of Desmoncus tested, there is an increase in Estr Province, South-west China, during two periods from towards the base of the axis (Isnard, Speck & Rowe, September to January 2003 and 2004. The climate is 2005; Rowe et al., 2004) and this is more similar to the dominated by the south-west monsoon bringing high pattern of change observed in arborescent, self- rainfall from May to October. supporting palms (Rich, 1986, 1987). In one species of Morphological characteristics of the plant stem Desmoncus, loss of the leaf sheath and a resultant loss were observed in the field and measured before the of rigidity is offset by an increase in Estr of the stem. mechanical tests. They include: internode lengths The fact that climbing stems of Desmoncus are not along main axes and branches; positions of buds and especially compliant, instead possessing quite stiff branching, and the state (living or dead) of apical mer- mechanical properties, suggests that there are rather istems and buds. different developmental constraints in climbing archi- tecture within the Arecoideae compared with the Calamoideae, or at least between Desmoncus and spe- MECHANICAL MEASUREMENTS cies belonging to the subtribe Calaminae. This study Young individuals were identifieded by self- on the bending mechanical properties of Plectocomia supporting or unstable branches with a leaf sheath himalayana Griff. (Calamoideae, Plectocomiinae) firmly encircling the stem along the entire length of aims to find out whether the mechanical architecture the axis. Old individuals were identified by climbing described for species of Calamus and Daemonorops axes where the leaf sheath has been lost from a large characterizes other rattan genera of the subfamily, or portion of the axis. Mechanical tests were carried out whether mechanical architecture and climbing strat- on four young individuals and on three older and egies are more diverse. longer climbing axes. Senescence and fragmentation Plectocomia himalayana has a climbing habit and of the leaf sheath were considered integral parts of is remarkable for growing in the Himalayas at alti- axis development and axes with leaf sheaths that tudes up to 2000 m. It has been reported in north-east were senescent or partially so were tested in bending. India, Laos and South Yunnan Province, China Selected axes were cut at the base, just above contact (Evans, 2001). The species grows naturally in the with the main stem (branches) or the rhizome (entire vicinity of Mengsong village, Xishuangbanna (Yun- ramets). The axis was then carefully stripped of nan), at an elevation of 1600 m where this study was leaves before detaching from the surrounding vegeta- carried out. A striking feature of P. himalayana is the tion and stored in humid conditions before mechani- formation of bulb-like shoots in the proximal part of cal tests. lower internodes, a feature that has been described The bending test protocol followed that of recent for the genus in P. elongata Mart. ex Blume (Fisher & analyses for 3-point bending (Rowe & Speck, 1996; Dransfield, 1979; Uhl & Dransfield, 1987). The ability Isnard, Rowe & Speck, 2003b) where consecutive to branch naturally from the aerial stem and form weights are added to a pannier suspended in the exact aerial adventitious roots is rare in climbing palms centre of the stem segment, which is supported in a © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 151, 83–91 MECHANICAL PROPERTIES AND BRANCHING IN PLECTOCOMIA HIMALAYANA 85 RESULTS MORPHOLOGY AND DEVELOPMENT OF THE CLIMBING HABIT Stems of Plectocomia himalayana are clustering and LS1 can reach at least 40 m in length and probably more. Cane diameters range between 1.5–3 cm without leaf LS2 sheaths and 2–5 cm with leaf sheaths (10 cm maxi- mum recorded, cf. Evans, 2001). The species is hapax- stem anthic, meaning that the shoots die after flowering
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