Vol.:(0123456789) 1 3
Oecologia
https://doi.org/10.1007/s00442-019-04452-7
PHYSIOLOGICAL ECOLOGY – ORIGINAL RESEARCH
Scaling of stem diameter and height allometry in 14 neotropical palm
species of diferent forest strata
Gerardo Avalos
1,2
· Maga Gei
1
· Luis Diego Ríos
3
· Mauricio Fernández Otárola
1
· Milena Cambronero
1
·
Carolina Alvarez‑Vergnani
1
· Olivia Sylvester
4
· Gustavo Rojas
1
Received: 9 November 2018 / Accepted: 27 June 2019
© Springer-Verlag GmbH Germany, part of Springer Nature 2019
Abstract
Tropical palms reach tree-like heights without a vascular cambium through sustained cell expansion and lignifcation of
primary tissues, but only a fraction of palms have been explored in their allometric relationships. Here, our main question
was to determine how palms depart from the traditional mechanical models developed for trees and how they approach the
theoretical buckling limit. We analyzed the stem allometry of 1603 palms of 14 species from diferent strata at 10 sites in
Costa Rica and Peru. We measured their ft to the stress, elastic, and geometric similarity models, and their position relative
to the maximum theoretical buckling limit calculated for trees. We evaluated the slope of the linear and logarithmic regres-
sions between stem diameter and height using logarithmic least squares, and standardized major axis regression (SMA),
expecting segregation according to canopy position and geographic location. Seventeen out of 19 statistically signifcant
models had SMA slopes > 1, and 11 had SMA slopes ≥ 2, departing from traditional mechanical models developed for trees.
Many species varied their allometry relative to geographic location. Canopy palms showed the highest regression ft but
had less steep slopes than understory and subcanopy species. Subcanopy and understory species were more underbuilt than
canopy palms, increasing height faster than diameter. Some of the tallest canopy palms surpassed the maximum buckling
limit whereas subcanopy and understory species were consistently below the buckling limit of record-size trees. Palm stem
allometry changed in response to environmental conditions.
Keywords Buckling limit · Mechanical stability · Palm architecture
Introduction
The palm family (Arecaceae) is one the most diverse, widely
distributed, and utilized groups of monocots, with nearly
2400 species in approximately 200 genera (Dransfeld et al.
2008). This high level of species diversity is refected in
varied architectural patterns, morphological structure, and
regeneration and life history strategies (Kahn and De Gran-
ville 2012). Palms develop very strong stems comparable
in magnitude to those of trees without producing woody
tissues generated by a vascular cambium. Instead, they
undergo sustained primary growth, which consists of the
division, expansion, and lignifcation of parenchyma cells
[the “difuse secondary thickening” of Schoute (1912) and
Waterhouse and Quinn (1978)] and can increase the cell
wall thickness of vascular fbers (Rich et al. 1986; Rich
1987b; Niklas 1993; Tomlinson et al. 2011). These rein-
forced primary tissues are concentrated close to the stem
periphery and around vascular bundles, and in many spe-
cies, their density decreases from the base toward the apex
of the stem and from the stem periphery toward the core of
the stem (Killmann 1983; Tomlinson et al. 2011). Ligni-
fed tissues result in increased resistance to biomechanical
strains, and thus, facilitate attaining considerable heights.
Communicated by Fernando Valladares.
* Gerardo Avalos
gerardo.avalos@ucr.ac.cr
1
Escuela de Biología, Universidad de Costa Rica, San Pedro,
San José 11501-2060, Costa Rica
2
The School for Field Studies, Center for Sustainable
Development Studies, 100 Cummings Center, Suite 534G,
Beverly, MA 01915, USA
3
Department of Biology, Indiana University, 1001 East Third
Street, Bloomington, IN 47405, USA
4
Department of Environment and Development, University
for Peace, Ciudad Colón, San José, Costa Rica