- Spatial pattern and process in forest stands within the Virginia piedmont - 37
Journal of Vegetation Science 16: 37-48, 2005
© IAVS; Opulus Press Uppsala.
Abstract
Question: Underlying ecological processes have often been
inferred from the analysis of spatial patterns in ecosystems.
Using an individual-based model, we evaluate whether basic
assumptions of species’ life-history, drought-susceptibility,
and shade tolerance generate dynamics that replicate patterns
between and within forest stands.
Location: Virginia piedmont, USA.
Method: Model verification examines the transition in forest
composition and stand structure between mesic, intermediate
and xeric sites. At each site, tree location, diameter, and status
were recorded in square plots ranging from 0.25 to 1.0 ha.
Model validation examines the simulated spatial pattern of
individual trees at scales of 1-25 m within each forest site
using a univariate Ripley’s K function.
Results: 7512 live and dead trees were surveyed across all
sites. All sites exhibit a consistent, significant shift in pattern
for live trees by size, progressing from a clumped understorey
(trees ≥ 0.1 m in diameter) to a uniform overstorey (trees >
0.25 m). Simulation results reflect not only the general shift in
pattern of trees at appropriate scales within sites, but also the
general transition in species composition and stand structure
between sites.
Conclusions: This shift has been observed in other forest
ecosystems and interpreted as a result of competition; how-
ever, this hypothesis has seldom been evaluated using simula-
tion models. These results support the hypothesis that forest
pattern in the Virginia piedmont results from competition in-
volving species’ life-history attributes driven by soil moisture
availability between sites and light availability within sites.
Keywords: Individual-based model; MUSE; Point pattern
analysis; Resource competition; Ripley’s K function; Soil
moisture.
Nomenclature: Harlow et al. (1996).
Abbreviations: ALB = Above-ground live biomass; BA =
Basal area; CSR = Complete spatial randomness; DBH =
Diameter at breast height; LAI = Leaf area index; MUSE =
Multistrata spatially explicit model; PET = Potential evapo-
transpiration.
Spatial pattern and process in forest stands
within the Virginia piedmont
Druckenbrod, Daniel L.
1*
; Shugart, Herman H.
2
& Davies, Ian
3
1
Environmental Sciences Division, Oak Ridge National Laboratory, P.O. Box 2008, Oak Ridge, TN 37831-6036, USA;
2
Department of Environmental Sciences, University of Virginia, 291 McCormick Rd., Charlottesville, VA 22904-4123, USA;
3
Ecosystems Dynamics Group, Research School of Biological Sciences, GPO Box 475, Canberra ACT 2601, Australia;
*
Corresponding author; Fax +18655768543; E-mail druckenbrodd@ornl.gov
Introduction
The piedmont forests of eastern North America have
a rich history of ecological research (Peet & Christensen
1987). Early research in the piedmont by Korstian &
Coile (1938), Kramer & Decker (1944), and Kozlowski
(1949) demonstrated the importance of competition for
light and soil moisture among individual trees in struc-
turing successional patterns in these forests. Similarly,
at a larger scale, Oosting (1942) characterized three
climax piedmont forest types along a moisture gradient
from mesic to xeric: Quercus alba - Q. velutina - Q.
rubra, Q. alba, and Q. alba - Q. stellata. These re-
sources, light and soil moisture, have recently been
shown to also influence the long-term temporal patterns
of individual tree growth in the piedmont (Druckenbrod
& Shugart 2004; Druckenbrod et al. 2003). Drawing
upon these insights, we examine the concurrent influ-
ence of light and soil moisture availability across scales
of individual trees to forest stands in the piedmont.
Using field survey data and individual-based modelling,
we evaluate whether competition for light and soil mois-
ture can reproduce piedmont forest patterns including
tree distributions within stands and forest composition
and structure among stands.
In a survey of individual tree spatial pattern in the
piedmont, Christensen (1977) reported a transition from
a clumped distribution of understorey trees to a random
or uniform distribution of overstorey trees. Field studies
of individual-tree spatial pattern over a range of other
forest types have generally revealed a similar transition
and have often suggested that competition for light
creates these spatial patterns (e.g. Cooper 1961; Laessle
1965; Whipple 1980; Good & Whipple 1982; Kenkel
1988; Moeur 1993; Ward et al. 1996; Busing 1998;
Mast & Veblen 1999; Van Pelt & Franklin 2000).
Watt (1947) first recognized the autogenic forma-
tion of spatial pattern in vegetation by observing a
landscape mosaic of regenerating forest patches. More
recently, ecologists have sought to explain this patch-