Ecological Modelling 154 (2002) 45 – 60
Growth dynamics of three tropical savanna grass species: an
individual-module model
Miguel F. Acevedo
a,
*, Josep Ravento ´s
b
a
Institute of Applied Sciences and Department of Geography, Uniersity of North Texas, P.O. Box 305279, Denton,
TX 76203, USA
b
Departamento de Ecologı ´a, Uniersidad de Alicante, Alicante, Spain
Received 2 August 2001; received in revised form 13 February 2002; accepted 4 March 2002
Abstract
We model the dynamics of grass plant growth as a collection of the individual dynamic behavior of shoots and
leaves. The model is inspired in data for plants of three species (Elyonurus adustus, Leptocoryphium lanatum and
Andropogon semiberbis ) of common grasses in the Venezuelan savannas that were sampled monthly for 1 year. These
species represent different architecture and regeneration response to fire. Modules (shoots and leaves) were counted
in each cell of a square grid in each one of several vertical levels. Module density per cell provides the horizontal
distribution within a level and is aggregated by level to obtain vertical distribution. Both distributions are simulated
by a dynamical model based on shoot emergence and mortality, elongation of shoots and leaves given by Richards’
equation, plus a few simple geometric considerations. For quantitative comparisons of model results to data, the
transient and final values for vertical distribution plus two metrics of horizontal distribution at each level, were
calculated for the simulation results and the field data. Proportion of occupied cells and maximum distance to the
center of growth were the two metrics selected to capture the dispersion and range of the horizontal distribution. The
model results indicate predictable final vertical profiles (of proportional density plus the two metrics) similar to the
profiles of measured distributions for each species. A reasonable prediction of the transient behavior was also
obtained but with larger deviations as evaluated by the root mean square error between model and data. Differences
in vertical and horizontal patterns of module density among species are explained by changing a set of parameter
values related to growth form and phenology. Thus, the model could be applied to generate plant functional types
for analysis of savanna dynamics subject to fire. With modifications, the model is potentially applicable to other grass
plants and other grassland ecosystems. © 2002 Elsevier Science B.V. All rights reserved.
Keywords: Growth; Phenology; Shoots; Richards’ equation; Savanna; Grasses; Venezuela; Fire
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1. Introduction
Sessile organisms, and especially terrestrial
plants, are discrete entities that live attached to
surfaces and interact mainly with their own or
* Corresponding author. Tel.: +1-940-565-2381; fax: +1-
940-565-4297.
E-mail address: acevedo@unt.edu (M.F. Acevedo).
0304-3800/02/$ - see front matter © 2002 Elsevier Science B.V. All rights reserved.
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