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Forest Ecology and Management
journal homepage: www.elsevier.com/locate/foreco
Changes in tree structure, composition, and diversity of a mixed-dipterocarp
rainforest over a 40-year period
Sisira Ediriweera
a,
⁎
, Champika Bandara
a
, David J. Woodbury
b
, Xiangcheng Mi
c
,
I.A.U.N. Gunatilleke
d
, C.V.S. Gunatilleke
d
, Mark S. Ashton
b
a
Department of Science and Technology, Uva Wellassa University, Badulla, Sri Lanka
b
School of Forestry and Environmental Studies, Yale University, New Haven, CT 06511, USA
c
State Key Laboratory of Vegetation and Environmental Change, Institute of Botany, Chinese Academy of Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, China
d
Botany Department, University of Peradeniya, Peradeniya, Sri Lanka
ARTICLE INFO
Keywords:
Borneo
Climate change
Elevation
Forest dynamics
Forest productivity
Mesua spp.
Shorea spp.
South Asia
Sri Lanka
Tree diversity
Tree mortality
Tree recruitment
ABSTRACT
Little work has examined the spatial and temporal changes of a tropical rain forest for long time periods. Here,
we present an analysis of long-term plot data from a Sri Lankan Mixed-Dipterocarp forest (MDF). The plots were
established in 1978 at three different elevations (low - 335m, medium - 560 m, and high - 915 m). At each site
we measured all trees ≥ 10 cm DBH in 16 or 18 plots. The total area sampled amounted to 12.5 ha. We recorded
166 tree species, with each site comprising between 95 and 121 species. Stem densities and basal areas were
different among sites, and declined over the period from 642.8 stems ha
-1
and 41.8 m
2
ha
-1
for the 1978
census, to 496.2 stems ha
-1
and 35.2 m
2
ha
-1
for the 2018 census. The lowest elevation exhibited the highest
recruitment, mortality and turnover among sites but patterns changed and became less marked over time. Forest
wide tree recruitment was about 1% y
-1
, while mortality was twice as much averaging about 2% y
-1
. Four
families made up between 44% (1978 census) and 54% (2018 census) of the trees sampled. Two families –
Clusiaceae, Dipterocarpaceae contributed up to 33.4% and 37.3% respectively, depending upon site and year of
census. The five most common species represent between 31 and 54% of the basal area and between 25 and 66%
of the stem density depending upon site. Percentage dominance by the most common species increased over the
40-year time period. Though many trees showed some degree of spatial differentiation, canopy trees showed
greater site overlap in their distribution than understory and sub-canopy species. Our results provide an im-
proved picture of variations in MDF structure and composition across the ever-wet realms of equatorial south
and southeast Asia. We suggest that dominance in dipterocarps and the degree of closed canopy structure, as
measured by basal area, is influenced by time, elevation, the degree of topographic variability within elevation,
and topographic interactions with variabilities in climate (drought and windstorms). We propose higher standing
basal areas and super dominance of dipterocarps in MDF are linked to site, succession, and landform stability.
Dipterocarps increase in dominance with succession (time), with topography (ridges greater than valleys) and
with elevation (lowland < hill < lower montane).
1. Introduction
There is growing need for long term demographic studies of forests
in order to evaluate the dynamics of tropical trees (Condit 1995; Swaine
and Whitmore 1988; Zimmerman et al. 1994). Worldwide only a few
rain forests have been monitored for extensive periods of time such as
in Budongo, Uganda (Sheil et al. 2000), La Selva, Costa Rica (Lieberman
and Lieberman, 1987) and in Australasia (Solomon Islands – Burslem
et al. 1998, 2000; Queensland, Australia – Murphy et al. 2013); all of
which have records that are > 40-years old.
Mixed dipterocarp forest (MDF) is the major forest type that covers
most of southeast Asia (Malaysia, Indonesia, Thailand, Philippines,
Vietnam, Cambodia, Laos) and the wetter parts of south Asia (Western
Ghats, India; southwest Sri Lanka) (Ashton and Seidler, 2014). It
comprises the largest commercial timber resource on a per hectare basis
of any other tropical forest (Sist et al. 2003), primarily because of ca-
nopy dominant trees in a single plant family - the Dipterocarpaceae - for
which the forest is named (Ashton and Seidler, 2014). The longest term
studies for MDF in Asia exist in Peninsular Malaysia (Manokaran and
Kochummen, 1987; 1990) and Sarawak, Borneo (Ashton and Hall 1992;
https://doi.org/10.1016/j.foreco.2019.117764
Received 2 September 2019; Received in revised form 8 November 2019; Accepted 8 November 2019
⁎
Corresponding author.
E-mail address: sisira@uwu.ac.lk (S. Ediriweera).
Forest Ecology and Management 458 (2020) 117764
0378-1127/ © 2019 Elsevier B.V. All rights reserved.
T