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The Mechanical Stability of the World's Tallest Broadleaf Trees

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The Mechanical Stability of the World's Tallest Broadleaf Trees bioRxiv preprint doi: https://doi.org/10.1101/664292; this version posted June 10, 2019. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. 1 1 The mechanical stability of the world’s tallest broadleaf trees 2 T. Jackson1,2*, A. Shenkin1, N. Majalap3, J. bin Jami4, A. bin Sailim4, G. Reynolds4, D.A. Coomes2, C.J Chandler5, 3 D.S. Boyd5, A. Burt6, Phil Wilkes6,7, M. Disney6,7, Y. Malhi1 4 1 – Environmental Change Institute, School of Geography and the Environment, University of Oxford, OX1 3QY, UK 5 2 – Forest ecology and conservation group, Department of Plant Sciences, University of Cambridge, CB2 3EA, UK 6 3 – Phytochemistry Unit, Forest Research Centre, Jalan Sepilok, 90715 Sandakan, Sabah, Malaysia 7 4 – South East Asia Rainforest Research Partnership (SEARRP), Danum Valley Field Centre, 91112 Lahad Datu, Sabah, Malaysia 8 5 – School of Geography, University of Nottingham, Nottingham, NG7 2RD, UK 9 6 – Department of Geography, University College London, WC1E 6BT, UK 10 7 – NERC National Centre for Earth Observation (NCEO), Leicester, UK 11 12 *Corresponding author – [email protected] 13 Abstract 14 The factors that limit the maximum height of trees, whether ecophysiological or mechanical, are the 15 subject of longstanding debate. Here we examine the role of mechanical stability in limiting tree 16 height and focus on trees from the tallest tropical forests on Earth, in Sabah, Malaysian Borneo, 17 including the recently discovered tallest tropical tree, a 100.8 m Shorea faguetiana. We use 18 terrestrial laser scans, in situ strain gauge data and finite-element simulations to map the 19 architecture of tall broadleaf trees and monitor their response to wind loading. We demonstrate 20 that a tree’s risk of breaking due to gravity or self-weight decreases with tree height and is much 21 more strongly affected by tree architecture than by material properties. In contrast, wind damage 22 risk increases with tree height despite the larger diameters of tall trees, resulting in a U-shaped 23 curve of mechanical risk with tree height. The relative rarity of extreme wind speeds in north Borneo 24 may be the reason it is home to the tallest trees in the tropics. 25 Introduction 26 Tall trees are an essential store of carbon and an inspiration to the public and to ecologists alike 27 (Bastin et al., 2018; Lindenmayer and Laurance, 2016; Lutz et al., 2018). Recently, the world’s tallest 28 tropical tree, a 100.8 m Shorea faguetiana (named “Menara”), has been discovered in Sabah, 29 Malaysian Borneo (Shenkin et al., 2019). Trees over 80 m tall have long been recognised in 30 temperate regions - notably the conifer Sequoia sempervirens in California and the broadleaf 31 Eucalyptus regnans in Tasmania, but the discovery of such tall trees in the tropics is recent. Record- 32 sized tropical trees have also recently been discovered Africa (Hemp et al., 2017) and South America 33 (Gorgens et al. in review). bioRxiv preprint doi: https://doi.org/10.1101/664292; this version posted June 10, 2019. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. 2 1 2 Figure 1 – Tree height against trunk diameter measured at breast height (dbh), or above the buttress in 3 buttressed trees, based on the continental allometric equation for Asia (Feldpausch et al., 2011) with trees from 4 this study overlaid. Menara, the world’s tallest tropical tree is at the far right. 5 Tree height growth is driven by the intense competition for light in young forest stands (MacFarlane 6 and Kane, 2017) but why emergent trees continue to grow upwards once they have escaped 7 competition, and what determines the mean and maximum height of forest canopies, is less well 8 understood. Why do the world’s tallest broadleaf trees grow to about 100 m in height, and not 50 m 9 or 500 m? 10 Possible answers to this question are that tree height is limited by hydraulics, carbohydrate supply or 11 by mechanics. The hydraulic limitation theory posits that the negative water potentials in the 12 vascular systems at the tops of trees, driven by gravity and by demand outstripping supply around 13 midday, inhibits the transfer of water from the vascular system into the cytoplasm of plants (Ishii et 14 al., 2014; Koch et al., 2004). This effect, and the increased risk of embolism, is predicted to lead to 15 lower tree heights in drier regions and is supported by the correlation between canopy height and 16 water availability at the global scale (Klein et al., 2015; Moles et al., 2009a). The carbohydrate 17 transport theory posits that it is the ability of leaves to supply carbohydrates to distant tissues 18 against resistance that limits tree to around 100 m (Jensen and Zwieniecki, 2013), and explains the 19 relationship between water availability and tree height as being determined by climate constraints 20 on leaf size. The mechanical limitation theory suggests that tree height is limited by either gravity, 21 on which most of the literature has focussed due to its tractability, or wind damage risk, which is 22 addressed in the current study. Givnish et al., (2014) showed that the cost of producing and 23 sustaining support tissues increases disproportionately with tree height, meaning that the risk of 24 mechanical damage will also increase. The relative importance of mechanical, hydraulic and 25 carbohydrate constraints will vary according to local climate, and the tallest broadleaf trees are likely bioRxiv preprint doi: https://doi.org/10.1101/664292; this version posted June 10, 2019. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. 3 1 to be found in regions with little seasonal water stress, such as aseasonal tropical (Klein et al., 2015; 2 Moles et al., 2009b). 3 In this study we focus on the mechanical limitation theory in the context of the tallest tropical 4 forests on earth, in northern Borneo. Our key questions are: 5 1. How does gravitational stability vary with tree height, architecture and material 6 properties? Terrestrial laser scanning (TLS) data allow us to estimate the weight of the 7 crowns of tall trees and so test the relative importance of variations in crown size and 8 material properties in terms of their effects on gravitational stability. 9 2. How does the risk of wind damage change with tree height? Taller trees are exposed to 10 higher wind speeds, but they also have better resistance to bending due to their higher trunk 11 diameter and modulus of elasticity. We use field measurements of strain under wind to 12 explicitly test whether these effects balance out. 13 3. Is Menara, the world’s tallest tropical tree, close to its mechanical limits? We focus 14 particular attention on Danum Valley in Sabah, home to Menara and many other tall trees, to 15 illustrate and explore the mechanical limits to tree height. 16 Results 17 Term Definition Gravitational max height The tallest a tree can grow without collapsing under gravity Modulus of elasticity A measure of a material’s resistance to bending K The ratio of crown weight to trunk weight Bending strain Extension per unit length – in this case along a tree trunk The slope of the best fit line relating bending strain to squared wind Wind-strain gradient speed The measured height of the tree divided by the maximum height Risk factor consistent with safety, either from wind or gravity 18 19 Table 1 – Definitions of key terms 20 Gravitational risk factors decrease with tree height within a forest canopy 21 We used two models to calculate the gravitational risk factors, the ratio of measured height to 22 theoretical maximum height each tree could reach before buckling under its own weight. The Euler 23 model (equation 1) represents a tree as a tapering cone with realistic trunk diameter and material 24 properties (Greenhill, 1881). The ‘top-weight’ model corresponds to a tapering cone with a top- 25 weight to represent the crown (King and Loucks, 1978). A novel aspect of this study is the inclusion bioRxiv preprint doi: https://doi.org/10.1101/664292; this version posted June 10, 2019. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. 4 1 of detailed 3D tree architecture based on TLS data. This allowed us to estimate the woody volume in 2 different parts of the tree and so estimate the weight of the stem and the crown separately. 3 Both models predicted a decrease in gravitational risk factors with tree height (Figure 2a). This 4 means that, as trees get taller, their radial growth is more than sufficient to compensate for their 5 increased height. We also calculated gravitational risk factors directly from continental height- 6 diameter allometric equations (Feldpausch et al., 2011) and found the same trend (Figure 2b). The 7 intercept and exact shape of these risk factors depend on the relationship between tree height and 8 K, the ratio of crown weight to stem weight, but the negative trend and difference between 9 continents do not (S5).
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