Forest Ecology and Management 171 (2002) 261–274

Phenological differences in tree water use and the timing of

tropical forest inventories: conclusions from patterns
of dry season diameter change
T.R. Bakera,*, K. Affum-Baffoea,b, D.F.R.P. Burslema, M.D. Swainea
a
Department of Plant and Soil Science, St. Machar Drive, University of Aberdeen, Aberdeen AB24 3UU, UK
b
Forest Management Support Centre, P.O. Box 1415, Kumasi, Ghana
Received 20 November 2000; received in revised form 5 October 2001; accepted 10 October 2001

Abstract

Interspecific variation in water-induced fluctuations in stem girth demonstrates the mechanisms promoting coexistence in
seasonally dry tropical forest. In addition, these fluctuations are a potential, but unevaluated, source of bias in measurements of
annual tree growth rates. To examine diurnal and seasonal patterns of stem diameter change, tree girth was measured over 2 years
(1997–1999), using dendrometer bands, for three species (Celtis mildbraedii, C. zenkeri and Strombosia glaucescens) in semi-
deciduous forest in Ghana. Soil matric potential was measured concurrently at 15 cm depth. In addition, measurements of all
trees >20 cm dbh on three, 1 ha plots were made at the beginning and middle of the 1998/1999 dry season. During the severe
1997/1998 dry season, soil matric potential declined below
1.5 MPa and two species showed significant stem shrinkage. For
the evergreen species, C. mildbraedii, there was a significant positive effect of tree diameter on stem shrinkage, and shrinkage
was greater in the second, compared to the first, half of the dry season. For the deciduous species, C. zenkeri, shrinkage was
reduced during the second half of the dry season, following leaf fall. During 1998/1999, soil matric potential, did not decline
below
1.5 MPa, and rates of girth change remained positive for all species. There were no significant effects of size or
phenology on the rate of girth change in the plot-based study. Deviations in annual increment calculated over successive monthly
intervals indicate that a 10-fold difference in soil water availability between measurement occasions can lead to a 4% bias in
estimates of annual growth. Measurements of forest plots should be made when inter-annual variation in soil water availability is
low. In this forest, measurements should, therefore, be made during the wet season, contrary to published recommendations.
# 2002 Elsevier Science B.V. All rights reserved.

Keywords: Dendrometer bands; Ghana; Permanent plots; Tree growth

1. Introduction division, and shorter term fluctuations caused by

Changes in tree diameter are a combination of two
factors: the increase in dry matter from cambial cell
* seasonal tropical climates, where soil water availabil-
Corresponding author. Present address: School of Geography,
University of Leeds, Woodhouse Lane, Leeds LS2 9JT, UK.
Tel.: þ44-113-233-3361; fax: þ44-113-233-3308.
E-mail address: t.baker@geog.leeds.ac.uk (T.R. Baker).
changes in tree water status (Kozlowski and Winget,
1964; Sheil, 1995). These short term fluctuations
occur over both diurnal and seasonal timescales.
For example, seasonal fluctuations are significant in
ity is periodically low, and tree diameter may decrease
in the dry season (Boaler, 1963; Daubenmire, 1973;
Lieberman, 1982; Reich and Borchert, 1984; Swaine

0378-1127/02/$ – see front matter # 2002 Elsevier Science B.V. All rights reserved.
PII: S 0 3 7 8 - 1 1 2 7 ( 0 1 ) 0 0 7 8 7 - 3
262 T.R. Baker et al. / Forest Ecology and Management 171 (2002) 261–274
et al., 1990; Reich, 1995; Baker et al., unpublished).
Changes in tree diameter in response to tree water
status are closely related to changes in soil water
availability, and are also influenced by tree water
use characteristics. Fluctuations in diameter have been
related to changes in stem water potential (Hinckley
and Bruckerhoff, 1975; Hinckley et al., 1978; Reich
and Borchert, 1982; Garnier and Berger, 1986), and to
the use of stored water (Reich and Borchert, 1984;
Holbrook, 1995; Reich, 1995). Studies of these water-
related changes in tree diameter in seasonal tropical
forests are important for two reasons. Firstly, they
allow hypotheses concerning interspecific differences
in water use to be tested, illustrating mechanisms
that may promote species coexistence. Secondly, they
allow us to test whether inter-annual variation in soil
water availability between years introduces a signi-
ficant bias in measurements of annual diameter
increment. Measurements of tropical forest plots are
increasingly being used to monitor long-term changes
in forest biomass at regional scales (Phillips et al.,
1998). Critiques of this work have focussed on arte-
facts introduced by the methodology itself (Clark, in
press). It is, therefore, important to quantify all the
potential errors and biases in forest inventory proce-
dures (Phillips et al., in press).
One of the major axes of variation allowing species
coexistence in seasonal tropical forests relates to water
use. The different strategies are exemplified by decid-
uous and evergreen species (Sobrado, 1986, 1993;
Fanjul and Barajas, 1987; Borchert, 1994; Eamus,
1999). In a dry forest in Venezuela, evergreen species
had smaller seasonal changes in leaf water potential
and wood water content than deciduous species
(Sobrado, 1986, 1993), and in a semi-deciduous forest
in Costa Rica dry season stem water potentials
declined to
2.5 MPa in evergreen species, compared
to <
5 MPa for deciduous species (Borchert et al., in
press). At the same site in Costa Rica, patterns of dry
season diameter change correlate with variation in
phenology (Daubenmire, 1973; Reich and Borchert,
1984; Reich, 1995). Deciduous species decrease in
girth substantially as soil water availability declines,
whereas evergreen species maintain positive girth
increment throughout the dry season. The physiolo-
gical studies suggest that the greater contraction of
stem diameter in deciduous species is caused by both
more negative plant water potentials, and greater use
of stored water. However, it is also possible that other
factors that influence tree water relations, such as tree
size and topographic position, may differ between the
populations that are being compared.
Tree size has a significant effect on water relations,
although the direction of that effect remains conten-
tious (Goldstein et al., 1998; Eamus, 1999; Meinzer
et al., 1999). Some studies indicate that greater tree
size is associated with improved dry season tree
water status. For example, large trees maintained
maximum rates of transpiration for longer during
the day than small trees in a seasonal forest in
Panama (Goldstein et al., 1998), and taller saplings
maintained higher pre-dawn water potentials during
the dry season in an Australian savanna (Eamus,
1999). However, in a seasonally dry forest in
Panama, small trees have been shown to use deeper
sources of soil water during the dry season than large
trees (Meinzer et al., 1999), suggesting that in forest
ecosystems, large trees experience more frequent and
severe periods of low soil water availability. To
attribute differences in dry season diameter change
to differences in phenology, we need to adequately
account for variation in tree size. Similarly, we need
to account for differences in topographic position,
as topographic differences in soil water availability
have been shown to cause differences in plant water
potential in a seasonally dry forest in Panama (Becker
et al., 1988).
The possibility that water-induced fluctuations in
stem diameter are a significant bias in longer term
measures of growth was suggested following a study
of long-term growth rates in moist forest in Uganda
(Sheil, 1997). Most stems showed a reduction in
growth rates when measured over wetter seasonal
periods, possibly as a result of field measurements
at these times being made only on bright, sunny days
when diurnal stem shrinkage is high (Sheil, 1997).
However, no previous studies have measured changes
in tree girth at a range of timescales, in order to
directly evaluate whether short-term fluctuations
can significantly bias longer term measurements of
growth. Repeated measurements of forest plots, car-
ried out over whole year intervals, assume that tree
water status is identical on the first and all subsequent
measurement occasions. However, this assumption is
unlikely to be correct. If tree water status is higher
on the second compared to the first measurement
 T.R. Baker et al. / Forest Ecology and Management 171 (2002) 261–274
occasion, then diameter increment will be overesti-
mated, as increased hydration will add to the annual
increment. If tree water status is lower, then annual
increment will be underestimated.
By studying diurnal and seasonal changes in tree
girth and soil water availability in semi-deciduous
forest in Ghana 1997–1999, this paper tests two
hypotheses:
1. Dry season stem shrinkage, and hence water
stress, is greater in deciduous compared to ever-
green species, having accounted for differences in
tree size and topographic position.
2. A significant bias is introduced into measurements
of annual increment by inter-annual variation in
soil water availability.
2. Methods
2.1. Study site and species
The study site was Tinte Bepo Forest Reserve
(78040 N, 28060 W) which is classified as moist semi-
deciduous forest (Hall and Swaine, 1981). Mean
annual rainfall, 1971–1993, recorded at the nearest
Ghana Meteorological Service station at Bechem,
15 km northeast of the site, is 1288 mm. There is a
strong dry season from December to March, when soil
matric potential, at 20–60 cm depth, may decline
below
1.5 MPa, and a less severe drier period from
July to September (Veenendaal et al., 1996). The site is
at approximately 300 m a.s.l., and in the study area
overlies fine grained, non-micaceous hornblende gran-
ite (Adu, 1993). The topography is undulating with an
elevational difference of 30–50 m between summits
and valleys, over horizontal distances of 300–500 m.
Maximum slopes are approximately 208. Reddish
brown sandy clay soils are found on the summits,
yellowish brown sandy clay soils on the slopes and
yellowish brown sandy clay loams and sandy loams in
the valleys (Adu, 1993).
There are approximately equal proportions of
individuals of evergreen and deciduous species in
semi-deciduous forest (Hall and Swaine, 1981). Com-
mon canopy species, which may exceed 60 m in
height, are Triplochiton scleroxylon (Sterculiaceae),
Celtis mildbraedii (Ulmaceae) and Nesogordonia
263
papaverifera (Sterculiaceae). Common understorey
species are Cleidion gabonicum (Euphorbiaceae),
Microdesmis puberula (Pandaceae) and Baphia nitida
(Papilionaceae).
The three principal study species were C. mildbraedii
Engl. (Ulmaceae), C. zenkeri Engl. and Strombosia
glaucescens J. Leonard (Olacaceae). C. mildbraedii is
an evergreen, non-pioneer species that is very common
in semi-deciduous forests in Ghana, found throughout
the Guineo-Congolian forest zone and in East African
coastal forests (Hawthorne, 1995). Its abundance
declines markedly as the landscape becomes wetter,
at both regional and local scales (Fig. 1(a), Swaine and
Hall, 1986; Hawthorne, 1995). It is an important fuel
for domestic use and making pestles (N. Gyakari,
pers. comm.). C. zenkeri is a deciduous, non-pioneer
species found across the Guineo-Congolian forest
zone (Hawthorne, 1995). In Ghana, its distribution is
biased towards dry, fertile sites (Fig. 1(b), Swaine,
1996). S. glaucescens is a very common evergreen,
non-pioneer species in Ghana and found throughout the
Guineo-Congolian region. Its distribution in Ghana is
biased toward wet, infertile sites (Fig. 1(c), Swaine,
1996). It is currently important for making telegraph
poles (N. Gyakari, pers. comm.).
2.2. Topographic definitions
Within areas of uniform soil parent material, the
water regime is an important determinant of colour in
tropical soils due to its influence on the content and
form of the predominant iron oxide present. Haemi-
tite, which gives the soil a bright red colour, is the form
typical of well-drained areas whilst the formation of
goethite is favoured under wetter conditions (Schwert-
mann and Taylor, 1989). As the particular focus of this
study was to examine the influence of topographic
variation on water supply, soil colour was used to
define landscape position (cf. Clark et al., 1998). Soil
colour was assessed using a Munsell Soil Colour
chart at 15 cm depth, 2 m away on the western side
of each selected tree (see below). There were clear
and consistent changes in soil colour downslope,
and different landscape positions were defined on
the basis of the hue (Oyama and Takehara, 1991).
Summit soils were red, hue 5 YR, slope soils yellow/
red, hue 7.5 YR and valley soils pale brown/grey,
hue 10 YR.
264 T.R. Baker et al. / Forest Ecology and Management 171 (2002) 261–274

2.3. Dendrometer bands
Adult trees greater than 20 cm dbh whose crowns
received at least some direct side light, i.e. Dawkins
crown classification of 4 or 5 (Dawkins, 1958), were
selected for study, over an area of approximately
12 ha, over several watersheds. The large study area
indicates that the study trees were typically well
dispersed (>20 m) and were considered independent
data points for the purposes of statistical analysis.
Seasonal changes in tree girth were monitored using
dendrometer bands constructed from 20 mm width,

Fig. 1. Distributions of: (a) C. mildbraedii, (b) C. zenkeri and (c) S. glaucescens in Ghana. The location of Tinte Bepo Forest Reserve, 250 mm
isohyets and extent of the forest zone (shaded area) are also shown.
 T.R. Baker et al. / Forest Ecology and Management 171 (2002) 261–274
Fig. 1. (Continued ).
150 mm thickness, hard-tempered aluminium and held
in place with a stainless steel spring. Shoe eyelets were
used to strengthen the holes where the spring held the
band. Measurements to 0.01 mm were made using
Vernier calipers of the distance between scribe marks
on the overlapping front and rear ends of the band.
Bands were placed at 1.3 m height or 50 cm above the
top of any fluting or buttress. Trunk climbers were
cut prior to fixing the band. From 23 July 1997 to 6
August 1997, bands were applied to 28 C. mildbraedii,
five C. zenkeri and three S. glaucescens trees; one
S. glaucescens was added to the sample on 22 August
1997, two C. mildbraedii on 1 October 1997 and two
C. zenkeri and one S. glaucescens on 17 October 1997.
The total sample sizes were, therefore, 30, 7 and 5 for
the three species, respectively, which reflects the
species relative abundance at the landscape scale in
this forest type.
Measurements of tree girth were made at approxi-
mately monthly intervals until May 1999. In addition,
on four occasions (18 September 1997, 11 October
1997, 22 October 1997, and 21 November 1997)
measurements were made of diurnal fluctuations for
C. mildbraedii, by comparing tree girth between
morning (08:00–11:00 h) and late afternoon (15:30–
17:30 h) measurements. Weekly measurements at the
265
start of the study indicated that a short period was
required for the bands to settle (Keeland and Sharitz,
1993). The results are not influenced by long-term
settling of the bands, as a control band placed on one
individual of C. mildbraedii that had a badly damaged
crown, showed no change in girth over the 2 years
of the study. Therefore, to obtain reliable readings,
those taken less than 30 days following the attachment
of a band were discarded (cf. Pélissier and Pascal,
2000). On seven occasions, out of a total of 837
measurements, slipped or damaged bands had to be
replaced. For calculations of the rate of girth change,
data from these individuals were discarded for at
least 30 days, and annual increments were only calcu-
lated for trees and periods with continuous, 12 month,
records.
2.4. Soil matric potential
Measurements of soil matric potential were made
concurrently with measurements of tree girth. The
filter paper method of Deka et al. (1995) was used
as it gives reliable results down to low matric poten-
tials of approximately
2 MPa, sampling is rapid and
it is inexpensive. A standard filter paper is equilibrated
over 6 days with a soil sample, the water content of the
266 T.R. Baker et al. / Forest Ecology and Management 171 (2002) 261–274
filter paper determined and then converted to matric
potential using calibration equations given by Deka
et al. (1995). From 11 October 1997, samples at 15 cm
depth were taken 2 m from the southern side of 14 C.
mildbraedii across the range of landscape positions.
On the second and subsequent measurement occa-
sions, samples were taken 50 cm from either side of
the original sample point, and then three sets of three
sample points, each 50 cm behind the previous set.
This systematic sampling pattern was then repeated,
moving clockwise, and assumes there is no systematic
variation in soil matric potential for a 3.5 m radius
around each tree. From 5 November 1997 until 8 April
1999, samples were taken at 15 cm depth around all
42 trees, at the same time as measurements of the
dendrometer bands.
2.5. Forest plot study
To test whether size or phenology-related patterns
of dry season stem shrinkage could be detected at the
stand level, measurements were also made of three
1 ha permanent sample plots (PSPs) in the same forest
reserve, during the 1998/1999 dry season. Measure-
ments were made at the start (November) and end
(January) of the dry season. The plots showed no
evidence of recent fire damage, and are part of the
on-going programme of permanent sample plot enu-
meration run by the Ghana Forest Service (Bird and
Sackey, 1991). A full account of the field procedures
are given in Wong (1997) and Affum-Baffoe (1999).
On each measurement occasion in this study, the
girth of each tree >20 cm dbh was recorded, at a
height of 1.3 m, or 50 cm above the top of any buttress
(diameter at reference height, drh).
2.6. Data analysis: tree water relations
The relationships between patterns of dry season
stem diameter change, phenology, tree size and topo-
graphy were examined in two ways. Firstly, the effect
of these factors on dry season diameter change was
tested during the severe 1997/1998 dry season using
the dendrometer band data. Three repeated measures
ANOVA were performed, comparing the first and
second half of the dry season, 21 November 1997
to 14 January 1998, and 14 January 1998 to 18 March
1998, respectively.
1. Dry season diameter change of C. mildbraedii was
analysed with topographic position as a factor, and
tree diameter and annual increment (calculated 18
September 1997 to 18 September 1998) as
covariates.
2. Dry season diameter change of C. mildbraedii and
C. zenkeri were compared in summit positions
with species as a factor, and diameter and annual
diameter increment included as covariates.
3. Dry season diameter change of C. mildbraedii and
S. glaucescens were compared in slope positions
with species as a factor, and diameter and annual
diameter increment included as covariates.
Secondly, the plot-based study allowed the influ-
ence of the same factors on dry season diameter
change to be assessed at the stand level. Trees were
grouped into size and phenology classes. The median
diameter was used as a threshold in a two-way classi-
fication of tree size, with individuals less than or
equal to 30.5 cm diameter classified as small. Species
were classified as deciduous or evergreen, following
Hawthorne (1995); species with variable vegetative
phenology, such as Cylicodiscus gabonensis (Mimo-
saceae), were omitted from the analysis. The resulting
dataset comprised 465 trees in four categories. As the
residuals from parametric analysis were not normally
distributed, differences between groups were tested
for using Kruskal–Wallis tests (Sokal and Rohlf,
1995).
2.7. Data analysis: bias in estimates of tree growth
The effect of inter-annual variation in soil water
availability on measurements of annual increment was
examined for individuals of C. mildbraedii growing
in summit positions. Annual increments were calcu-
lated from successive measurement occasions, with
values for exact years obtained by linear interpolation
between measurement dates in the second year of
study. To remove the influence of temporal change
in actual growth over the course of the study, the data
were detrended using linear regression. The residuals
of this regression were then compared with the change
in mean soil matric potential between years. Similar
to the calculation of successive annual increments,
exact values for soil matric potential on equivalent
dates during the second year of study were calculated
 T.R. Baker et al. / Forest Ecology and Management 171 (2002) 261–274 267

by linear interpolation of mean logged soil matric
potential data.
3. Results
3.1. Soil matric potential regime
The soil matric potential at 15 cm depth was higher
in valley than slope and summit positions, apart from
during the late dry season of 1997/1998, when all sites
became very dry, and periodically during the wet
season when water availability was similarly high
in all sites (Fig. 2). Less negative minimum soil matric
potentials during November 1998 to March 1999,
indicate that the dry season was less severe during
the second year of the study (Fig. 2).
3.2. Magnitude of fluctuations
No significant diurnal girth fluctuations were found
for C. mildbraedii, either on the four occasions when
they were measured, or when the data were combined
(Table 1). However, significant dry season stem
shrinkage occurred for two of the three species, and
for C. mildbraedii, the magnitude of the mean dry

Fig. 2. Rates of change of girth for C. mildbraedii in summit and valley positions (slope position not shown), C. zenkeri and S. glaucescens,
and the soil matric potential regime in summit, slope and valley positions, at 15 cm depth, in semi-deciduous forest in Ghana, from October
1997 to April 1999. For the rate of girth change, significant differences between species, tested using univariate F-tests, are indicated,

p < 0:05, p < 0:01, p < 0:005.
268 T.R. Baker et al. / Forest Ecology and Management 171 (2002) 261–274

Table 1
Diurnal and dry season stem girth change in C. mildbraedii, C. zenkeri and S. glaucescens in semi-deciduous forest in Ghana, 1997/1998a

Diurnal Dry season

Mean  S:E: n Significance Mean  S:E: % of % of annual n Significance

(mm) (mm) diameter increment

C. mildbraedii 0:02  0:02 59 ns
0:68  0:13 0.18 15.2 30 <0.005

C. zenkeri
1:28  0:34 0.51 17.6 7 <0.005
S. glaucescens
0:11  0:14 0.03 17.9 5 ns
a
Mean diurnal fluctuations are calculated from morning (08:00–11:00 h) and late afternoon (15:30–17:30 h) measurements made on 18
September 1997, 11 October 1997, 22 October 1997, and 21 November 1997. The dry season is defined as 21 November 1997 to 18 March
1998. Z-tests were used to test if changes in girth were significantly different from zero.

season change was more than 30 times greater than Table 2

mean daily fluctuation (Table 1). Table of significance of repeated measures ANOVA of the effects
of size, annual increment, size, topographic position and time (first
and second half of the dry season), on dry season diameter change
3.3. Dendrometer study: tree water relations of C. mildbraedii in semi-deciduous forest in Ghana, 1997/1998 (p-
values in bold are significant at p < 0:05)
For C. mildbraedii, tree diameter had a significant F p
main effect on the degree of dry season diameter
change, indicating that larger trees shrank more than Size 10.8 0.00
Annual increment 3.0 0.10
smaller trees during the severe 1997/1998 dry season
Position 3.2 0.06
(Fig. 3, Table 2). Annual diameter increment did Time 2.9 0.10
not have a significant main effect, but there was a Time  size 0.7 0.40
significant time/increment interaction (Table 2). Time  increment 7.5 0.01
Although trees better supplied with water in valley Time  position 2.0 0.15

Fig. 3. Dry season change in diameter, 1997/1998, as a function of tree diameter for C. mildbraedii in a semi-deciduous forest in Ghana.
Linear regression: y ¼ 0:33
0:03x, F ¼ 11:9, p < 0:005.
 T.R. Baker et al. / Forest Ecology and Management 171 (2002) 261–274 269

Fig. 4. Mean and standard error of change in tree girth for C. mildbraedii and C. zenkeri during the early and late dry season 1997/1998,
November–January and January–March, respectively, in semi-deciduous forest in Ghana. For C. zenkeri, leaf fall occurred in all trees between
January and March measurements.

positions tended to shrink less (Fig. 2), the effect of
topographic position on dry season diameter change
was marginally non-significant (Table 2).
Evergreen and deciduous species growing in the
same topographic position showed significant differ-
ences in dry season diameter change (Fig. 4, Table 3).
In the comparison of C. zenkeri and C. mildbraedii in
summit positions, there was a significant time/species
interaction (Table 3). The rate of stem shrinkage
increased in the evergreen species, C. mildbraedii,
as the dry season progressed, but decreased in the
deciduous species, C. zenkeri, following leaf fall,
which occurred between January and March 1998
measurements in all trees of this species (Fig. 4).
However, there were no differences between the two
evergreen species, C. mildbraedii and S. glaucescens,
when they were compared in slope positions (Table 3).
3.4. Plot-based study: tree water relations
There were no differences in median diameter
change between small and large, deciduous and

Table 3
Table of significance from repeated measures ANOVA of the effects of size, annual increment, size, species and time, on dry season diameter
change of an evergreen and deciduous species (C. mildbraedii and C. zenkeri) and two evergreen species (C. mildbraedii and S. glaucescens)
in equivalent topographic positions in semi-deciduous forest in Ghana, November–January and January–March, 1997/1998 (p-values in bold
are significant at p < 0:05)

C. mildbraedii/C. zenkeri C. mildbraedii/S. glaucescens

F p F p

Size 19.2 0.00 0.29 0.60

Annual increment 0.6 0.44 2.87 0.12
Species 13.4 0.00 0.60 0.46
Time 3.2 0.09 0.17 0.69
Time  size 1.5 0.24 1.49 0.25
Time  increment 14.3 0.00 1.57 0.24
Time  species 52.0 0.00 1.77 0.22
270 T.R. Baker et al. / Forest Ecology and Management 171 (2002) 261–274

Fig. 5. Successive mean annual increments, S.E., for C. mildbraedii in summit positions in semi-deciduous forest in Ghana, 1997/1999.
Annual increments are calculated from successive measurement occasions during the first year of study. Linear regression: y ¼ 11:8 þ 0:026x,
F ¼ 263, p < 0:005.

evergreen species, in the plot-based study, during the
1998/1999 dry season (H ¼ 2:73, d:f: ¼ 3, p ¼ 0:434,
overall mean diameter change ¼ 0:03  0:01 cm,
median change ¼ 0:00 cm). The effects of size and
phenology may only be important during particularly
dry years. In the dendrometer study, higher rates of
girth change during the 1998/1999 dry season com-
pared to 1997/1998, reflected the less severe dry
season and resulted in similar patterns in all three
species, in contrast to the previous year (Fig. 2).
3.5. Dendrometer study: hypothesis 2
As there was no evidence for significant diurnal
fluctuations in tree girth, only the impact of seasonal
changes on the measurement of annual increment was
evaluated further. Annual increments calculated from
successive measurement occasions show a steady
significant increase over the study (Fig. 5). However,
there is also substantial variation around this relation-
ship, particularly for annual increments initiated dur-
ing the course of the dry season (Fig. 5). There was a
significant positive correlation between the difference,
and mean, of soil matric potential measured in suc-
cessive years (Fig. 6). This result demonstrates that
soil matric potential was more variable at generally
dry times of year. In addition, the difference in soil
matric potential between years is correlated with the
residual error in the calculation of annual increment
(Fig. 7). The significant regression indicates that a 10-

Fig. 6. Relationship between the mean, and variation, in soil matric potential between 1997/1998 and 1998/1999. Linear regression:
logðyÞ ¼ 0:45 logðxÞ
0:013, F ¼ 9:6, p < 0:05.
 T.R. Baker et al. / Forest Ecology and Management 171 (2002) 261–274
Fig. 7. Residual error in linear increase of annual increment through time, plotted against the difference in soil matric potential between years.
Linear regression: y ¼
0:57x
0:31, F ¼ 5:1, p < 0:005.
fold increase in soil matric potential between mea-
surement occasions leads to an overestimate of annual
increment by 0.5 mm, or approximately 4%. In this
study, the most extreme errors in the calculation of
annual increment were found in measurements at the
height of the dry season, when annual growth is
overestimated by more than 9% (Fig. 5).
4. Discussion
4.1. Tree water relations
As dry season diameter change only represents an
indirect measure of tree water status, inferring species
and size effects on tree water relations from these data
must be done with caution. Seasonal decreases in stem
diameter are caused by water loss from elastic storage
elements such as the phloem, cambium and sapwood
parenchyma (Holbrook, 1995). Close correlations
have been obtained between xylem water potential
and stem diameter over diurnal timescales (Worrall,
1966; Hinckley and Bruckerhoff, 1975; Hinckley et al.,
1978; Reich and Borchert, 1982; Garnier and Berger,
1986), but extending this relationship over longer,
seasonal timescales is more complex. Specifically,
inferring species and size effects on seasonal changes
in plant water potential from patterns of stem diameter
change assumes that tissue capacitance does not vary
across species and size classes. Tissue capacitance,
271
defined as the change in water content per unit change
in water potential, per unit volume (Holbrook, 1995),
does, however, vary between trees, due to differences
in tissue structure or the relative proportions of dif-
ferent tissues. For instance, variation in bark thickness
is important (Borchert, 1994). As phloem is an elastic
tissue, absolute declines in stem diameter, for a given
reduction in plant water potential, will be greater in
trees with thicker bark. In addition, differences in
growth rate may be important, as fast growing trees
produce larger cells that may show a greater reduction
in volume for the same change in water potential. The
result of these effects is that a given reduction in water
content (stem diameter) in two trees, that are different
species or sizes, may not reflect similar changes in
plant water potential. In this study, the effect of
differences in growth rate can be discounted, as sig-
nificant effects of tree size and species on dry season
diameter change were found, independently of varia-
tion in annual increment (Table 2). However, differ-
ences in bark thickness are important for interpreting
the relationships between dry season diameter shrink-
age and tree water relations.
The greater shrinkage of larger trees of C. mild-
braedii during the 1997/1998 dry season is consistent
with the suggestion that large trees experience greater
water deficits than small trees during the dry season.
The greater transpirational demand of a larger canopy
apparently exceeds the ability of a larger root system
to supply water. Research on the sources of dry season
272 T.R. Baker et al. / Forest Ecology and Management 171 (2002) 261–274
water and patterns of tree mortality has also suggested
that large trees are more susceptible to seasonal
drought. For example, by relating the isotopic signa-
ture of xylem sap to groundwater values, large trees
were found to tap shallower sources of water than
small trees during the dry season in a seasonally dry
forest in Panama (Meinzer et al., 1999). In addition,
drought related mortality was 50% higher in trees
>60 cm dbh, compared to individuals <30 cm dbh,
following the 1983 drought in Borneo (Wirawan,
unpubl. rep., 1983, in Goldammer and Seibert, 1990).
However, the suggestion that large trees experience
greater seasonal water deficits than small trees should
be treated with caution. Although the increased stem
shrinkage of larger trees must result from greater
water loss from elastic storage tissues, this does not
necessarily reflect the development of more negative
plant water potentials. Bark thickness increases with
tree size in C. zenkeri in Ghana (Orgle, 1994); if the
same relationship occurs in its congener, the effect of
size on dry season diameter change in C. mildbraedii
may be, at least in part, due to changes in capacitance,
in addition to plant water potential.
Topographic differences in soil water availability
have been shown previously to cause differences in
plant water potential in a seasonally dry forest in
Panama (Becker et al., 1988). In this semi-deciduous
forest in Ghana, topographic position did not have a
significant independent effect on diameter change,
although trees in sites with higher soil water avail-
ability did tend to decrease in diameter less during the
severe 1997/1998 dry season (Fig. 2). However, soil
water availability was an important determinant of
inter-annual differences in dry season diameter
change. No differences were found between species
and size classes during the second year of the study in
either the dendrometer or plot-based study, and the
mean rate of girth change did not decline below zero
for any species (Fig. 2).
The greater shrinkage of C. zenkeri compared to C.
mildbraedii during the first half of the 1997/1998 dry
season, suggests that water loss was greater, and more
negative water potentials were developed, in the
deciduous compared to the evergreen species. These
patterns are consistent both with patterns of dry season
diameter change in a dry forest in Costa Rica (Reich
and Borchert, 1984; Borchert et al., in press), and the
greater seasonal fluctuations in leaf water potential
and wood water content found in deciduous, compared
to evergreen, species in a dry forest in Venezuela
(Sobrado, 1986, 1993). Leaf abscission was an effec-
tive strategy for conserving water in C. zenkeri, as the
rate of shrinkage dropped following leaf fall in the
second half of the 1997/1998 dry season (Fig. 4).
However, two factors may confound the comparison
of deciduous and evergreen species. Firstly, differ-
ences in bark thickness between these two species
were not measured, and the magnitude of diameter
change may not directly reflect differences in plant
water potential, as discussed above. Secondly, the
species may differ in rooting depth allowing differ-
ential access to dry season water. Rooting depth was
not studied here, but the forest occurs on deep soils
(Adu, 1993) and there was no evidence of the forma-
tion of a hard pan in soil pits dug to 1 m depth.
However, studies of the stable hydrogen isotope com-
position of xylem sap and groundwater in seasonally
dry forest in Panama have shown that evergreen
species have access to water at greater depth than
deciduous species (Jackson et al., 1995). It is possible
that either differences in bark thickness or rooting
depth mean that the patterns of dry season diameter
change do not reflect equivalent differences in water
stress, or are not influenced by leaf phenology. How-
ever, the significant species/time interaction related to
leaf fall in C. zenkeri (Table 2), provides strong
support for the hypothesis that dry season water
deficits are greater in the deciduous compared to
the evergreen species, and that it is caused by differ-
ences in leaf phenology.
4.2. Errors in forest plot measurements
No evidence was found here for significant diurnal
fluctuations in stem girth, and hence no evidence that
they can generate significant errors in measurements
of annual increment (Sheil, 1997). It is possible that
dendrometer bands are unable to detect very small
changes over this short timescale. In contrast, Sheil
(1997) cites a study where multiple measurements of
tree girth using a tape measure found significant
diurnal changes in stem diameter. Alternatively, the
shade tolerant species C. mildbraedii, may show little
change in diameter over these timescales, possibly as a
result of a low proportion of elastic storage tissues in
the trunk.
 T.R. Baker et al. / Forest Ecology and Management 171 (2002) 261–274
Significant water-related changes in stem diameter
do result in errors in estimates of growth. The steady
increase in growth measured over successive, over-
lapping, time periods is caused by the decreasing
contribution of the 1997/1998 dry season to the cal-
culation of annual increment. Rates of wet season
expansion of these trees remained constant between
years, whereas dry season diameter change was sig-
nificantly higher in the second year of study (Baker
et al., unpublished). The hypothesis that increased soil
water availability on successive measurement occa-
sions leads to an overestimate of growth is supported,
as the deviations in annual increment about this trend
are negatively correlated with the difference in soil
matric potential between years (Fig. 7). Although the
potential error is small, it will be important where
intercensus intervals are short, in forests with a
strongly seasonal climate, and where inventories take
several months to complete.
Published recommendations on methods of tropical
forest inventory suggest that the optimum time to
measure forest plots is during the dry season, when
inter-annual variation in water-induced changes in tree
diameter are assumed to be minimal (Sheil, 1995).
However, this study indicates that variation in soil
water availability is greatest during the dry season
(Fig. 6). Although this study only compares 2 years,
one of which is strongly influenced by an El Niño
event, it is likely that the patterns described here are
representative of West African forests, and will be
reflected over longer timescales. Climatic fluctuations
during El Niño events are likely to be the main
determinant of inter-annual variation in climate, and
in this region, El Niño events typically lead to more
severe dry seasons (Swaine et al., 1997). Less bias
would be introduced into estimates of growth if enu-
merations were carried out during the wet season, to
avoid the large inter-annual variation in dry season
patterns of stem diameter change, caused by variation
in dry season soil water availability. This study pro-
vides direct evidence to support a similar recommen-
dation made for plots in wet evergreen forests with a
monsoon climate in the Western Ghats of India (Pélis-
sier and Pascal, 2000). Finally, in large inventory
programmes that maintain plots in a variety of forest
types, with a full-time schedule of remeasurements,
care should be taken in the timing of plot enumeration.
Measurements of plots in the climatically wettest
273
forest type should be made during the driest time of
year, and of the driest forest plots at the wettest time of
year, to minimise the effect of inter-annual variation in
dry season soil water availability on estimates of growth.
Acknowledgements
This work was funded by a University of Aberdeen
Faculty Studentship (TB) and the Department for
International Development (TB/KAB). Yaw Atua-
hene, William Asante and Raymond Votere provided
invaluable assistance with the fieldwork, and Doug
Sheil and Rolf Borchert gave very helpful comments
on a previous version of the manuscript.
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