Vol. 6/1,2, pp.

21–36 Perspectives
© Urban & Fischer Verlag, 2003 in Plant Ecology,
http://www.urbanfischer.de/journals/ppees Evolution and
Systematics

Variation in tropical forest growth rates:

combined effects of functional group composition
and resource availability

Timothy R. Baker1,2,3,*, Michael D. Swaine1 & David F.R.P. Burslem1

1
School of Biological Sciences, University of Aberdeen, Aberdeen, UK
2
Max-Planck-Institut für Biogeochemie, Jena, Germany
3
Centre for Biodiversity and Conservation, School of Geography, University of Leeds, UK

Received: 12 February 2003 · Revised version accepted: 28 March 2003

Abstract
Rates of tree growth in tropical forests reflect variation in life history strategies, con-
tribute to the determination of species’ distributional limits, set limits to timber harvesting
and control the carbon balance of the stands. Here, we review the resources that limit tree
growth at different temporal and spatial scales, and the different growth rates and re-
sponses of functional groups defined on the basis of regeneration strategy, maximum size,
and species’ associations with particular edaphic and climatic conditions.
Variation in soil water availability determines intra- and inter-annual patterns of growth
within seasonal forests, whereas irradiance may have a more important role in aseasonal
forests. Nutrient supply limits growth rates in montane forests and may determine spatial
variation in growth of individual species in lowland forests. However, its role in determin-
ing spatial variation in stand-level growth rates is unclear. In terms of growth rate, we
propose a functional classification of tropical tree species which contrasts inherently fast-
growing, responsive species (pioneer, large-statured species), from slow-growing species
that are less responsive to increasing resource availability (shade-bearers, small-statured
species). In a semi-deciduous forest in Ghana, pioneers associated with high-rainfall
forests with less fertile soils, had significantly lower growth rates than pioneers that are
more abundant in low-rainfall forests with more fertile soils. These results match patterns
found in seedling trials and suggest for pioneers that species’ associations with particular
environmental conditions are useful indicators of maximum growth rate.
The effects of variation in resource availability and of inherent differences between species
on stand-level patterns of growth will not be independent if the functional group compo-
sition of tropical forests varies along resource gradients. We find that there is increasing
evidence of such spatial shifts at both small and large scales in tropical forests. Quantify-
ing these gradients is important for understanding spatial patterns in forest growth rates.

Key words: irradiance, maximum size, nutrient supply, pioneer, regeneration groups,
water availability

*Corresponding author: Centre for Biodiversity and Conservation, School of Geography, University of Leeds LS2 9JT, UK;
e-mail: t.baker@geog.leeds.ac.uk

1433-8319/03/6/01-02-021 $ 15.00/0
22 T. R. Baker et al.

Introduction important differences between tropical forests in the

An understanding of the patterns of tree growth is a
fundamental goal of ecological research in tropical
forests. Interspecific variation in maximum potential
growth rate consistently emerges as one of the most
important factors in the definition of robust functional
groups, as growth rate integrates numerous traits that
underlie trade-offs among strategies for resource ac-
quisition, defence against natural enemies, and alloca-
tion to reproduction. Such groups provide practical
and meaningful classifications of tropical forest
species, which are needed both by foresters, for mod-
elling growth and yield, and by ecologists, to explain
the life history diversity in tropical forest trees (Van-
clay 1994; Richards 1996; Whitmore 1998; Turner
2001). In addition to these inherent differences be-
tween species, growth also varies with resource avail-
ability. At the species level, this variation may help to
explain the limits to species distributions (e.g. Gu-
natilleke et al. 1996; Veenendaal et al. 1996a). At the
stand level, understanding the environmental factors
that control tropical forest productivity is critical for
quantifying the carbon balance of tropical forests.
Spatial variation in stand-level growth rates will de-
pend on both variation in resource availability and
any differences in the functional composition of tropi-
cal forests. The large variation in forest composition
and dynamics at both small and large scales (Phillips
et al. 1994, in press; Burslem & Whitmore 1999, in
press; ter Steege et al. 2000) suggests that there may be
relative abundance of different functional groups.
These differences may be as important as gradients in
resource availability for determining current and fu-
ture patterns of forest productivity.
Building on the writings of Budowski (1965), Tim
Whitmore made an important contribution both to the
description of tropical tree functional groups and to
highlighting their significance to central questions of
tropical forest ecology (Whitmore 1974, 1975, 1998).
He also drew attention to the relationship between
plant functional traits, including growth rate, and
tropical forest silviculture (Whitmore 1975). Whit-
more’s promotion of a fundamental dichotomy be-
tween fast-growing, gap-demanding tree species (la-
belled ‘pioneers’) and slow-growing, non-gap-deman-
ders (‘non-pioneers’ or ‘climax’ species) is, perhaps,
one of his best known and most frequently cited con-
tributions (Whitmore 1975, 1984, 1998; Swaine &
Whitmore 1988). This perspective was undoubtedly
influenced by Whitmore’s own research on the dynam-
ics and growth rates of seedlings, saplings and large
trees of the twelve most common large tree species
growing on his plots on Kolombangara in the
Solomon Islands during the period 1964–1971 (Whit-
more 1974). Based on an initial seven-year study of
seedling responses to canopy openings, Whitmore
(1974) classified these twelve species into four groups
(Table 1) and emphasized the importance of differ-
ences in growth rate between species as a determinant
of group membership. A link between the four groups

Table 1. Characteristics of the twelve species studied since 1964 in lowland tropical rain forest on Kolombangara, Solomon Islands. The four species’ groups
among the twelve common timber tree species are classified according to the conditions required for seedling establishment and onward growth and are
based on observations over 6.6 years over the interval 1964–1971 (Whitmore 1974). Nomenclature follows sources described in Burslem & Whitmore (1999).

Shade-tolerance Species Conditions Conditions Wood density

class to establish to grow up (kg m–3)1

I Dillenia salomonensis High forest High forest 550

Maranthes corymbosa High forest High forest 720
Parinari papuana ssp. salomonensis High forest High forest 660
Schizomeria serrata High forest High forest 490

II Calophyllum neo-ebudicum High forest or small gaps High forest/gaps 500

Calophyllum peekelii High forest High forest/gaps 480
Pometia pinnata High forest or disturbed High forest 590
or ?small gaps

III Campnosperma brevipetiolatum High forest or gaps Gaps 330

Elaeocarpus angustifolius High forest Gaps 350

IV Endospermum medullosum Mostly gaps Gaps 370

Gmelina moluccana Mostly gaps Gaps 410
Terminalia calamansanai High forest, soon dying Gaps 460
except in gaps
1
Data from Anonymous (1976, 1979).

Perspectives in Plant Ecology, Evolution and Systematics (2003) 6, 21–36


defined for the twelve species growing on Kolomban-
gara and the dichotomy proposed in his later writings
can be made if the three species in Group IV of Table 1
are treated as pioneers, and the remaining species as
non-pioneers.
Since the mid 1970s, there has been a proliferation
of research on the characteristics that define the main
functional groups of tropical trees and their relation-
ship to forest dynamics and regeneration, expanding
and extending the ideas of Whitmore and his predeces-
sors and contemporaries. Some authors have ques-
tioned the existence of a fundamental dichotomy of
life history types among tropical trees (e.g. Alvarez-
Buylla & Martinez-Ramos 1992; Grubb 1996), and
the issue has been comprehensively reviewed several
times (Brokaw 1985; Denslow 1987; Brown & Jen-
nings 1998; Brokaw & Busing 2000; Turner 2001;
Burslem & Swaine 2002). Here, we specifically exam-
ine how useful these concepts are in understanding
variation in tree growth rates over gradients in re-
source availability. We consider the following specific
questions:
1. What is the evidence for resource limitation of
tropical tree growth?
2. How do functional groups defined in various
ways differ in their growth responses to resource avail-
ability?
3. Are there gradients in the functional-group com-
position of tropical forests, and are they important in
determining variation in stand-level growth?
We consider each of the major resources (water and
irradiance) or group of resources (nutrients) known to
limit plant growth over spatial gradients, both sepa-
rately, and, where possible, in combination. We adopt
a broad definition of growth rate in order not to im-
pose severe limitations on the scope of the review, and
emphasize evidence from field-based studies of adult
trees. Thus we include studies of litter fall, fine-root
production, leaf-level photosynthesis, and phenology
as well as direct measures of trunk growth based on
measurements of diameter change or the width of an-
nual rings.
Factors limiting tree growth
Water
The total amount of rainfall is one of the most impor-
tant factors setting the limits to the distribution of
forests as opposed to woodland or thicket in the trop-
ics (Holdridge 1967; Walter 1979; White 1983; Wood-
ward 1987). However, the forested regions experience
considerable spatial and temporal variation in the
availability of soil water that acts as a major limiting
Variation in tropical forest growth rates 23
factor on overall rates and temporal patterns of
growth.
The importance of soil water availability is most ap-
parent within forests with strongly seasonal climates.
Over ten years in dry forest in Mexico (mean annual
rainfall 707 mm) the annual increment of two decidu-
ous species correlated with rainfall during the mid
wet-season (Bullock 1997). Within the same forest,
over a five-year period when annual rainfall ranged
from 485 to 1331 mm, Whigham et al. (1990) found
that the annual production of leaf litter correlated pos-
itively with annual rainfall and that the mean basal
area increment per tree correlated positively with total
rainfall during the previous two years. Soil water
availability also controls the timing of growth within
seasonal forests. For the evergreen species, Exostema
caribaeum, in dry forest in Puerto Rico (mean annual
rainfall 929 mm), Lugo et al. (1978) found maximum
daily rates of photosynthesis were five times higher
during the wet season compared to the dry season. In
addition, marked fluctuations in tree girth in parallel
with monthly rainfall were described by Swaine et al.
(1990) for very dry tropical forest in Ghana (c. 750 mm
yr–1), where seasonal variation in girth was about ten
times greater than the underlying annual increment in
Millettia thonningii. Direct measurement of wood pro-
duction in seasonal forest from examination of the
pattern of cambial activity, by measuring either the
width of the band of differentiating xylem which
stains for cellulose (Amobi 1973), or the width of un-
lignified xylem (Lowe 1968), has also demonstrated
the importance of increased soil water availability for
the initiation of growth in seasonal forests (Borchert
1999).
Soil water availability also influences the timing and
amount of growth in forest with substantially higher
mean annual rainfall than the previous examples. For
instance, in a semi-deciduous forest in Venezuela
(mean annual rainfall 1700 mm), using tree ring
chronologies of seven species, Worbes (1999) reported
positive correlations between annual rainfall and
mean annual growth rates, and at three sites in season-
al forest in Panama, Devall et al. (1995) found that an-
nual rainfall correlated with variation in tree ring
width for three species. In contrast to these correlative
studies, field experiments that directly test whether
water supply limits growth in a particular forest are
rare. However, an irrigation experiment over five dry
seasons in semi-evergreen forest in Panama (mean an-
nual rainfall 2600 mm) showed that ameliorating the
seasonal drought could increase fine-root production
and leaf longevity, and influence phenology. Dry sea-
son community-level fine-root production, measured
using in-growth cylinders with root-free soil, was five
times greater in irrigated plots than in control plots
Perspectives in Plant Ecology, Evolution and Systematics (2003) 6, 21–36
24 T. R. Baker et al.
(Cavelier et al. 1999), and dry season leaf fall was de-
layed for two out of nine tree species (Wright &
Cornejo 1990). In addition, for three shrub species,
the proportion of leaves retained, of those that were
newly flushed at the start of the experiment, was ap-
proximately 20% higher over the first two years in the
irrigated plots (Mulkey et al. 1993). However, this ex-
periment also demonstrated that not all aspects of
growth, and not all species, are limited by seasonal
water shortage in this forest. Irrigation did not affect
community level leaf-litter production (Cavelier et al.
1999), stem diameter growth of five shrub species
(Wright 1991), or the timing of leaf fall for seven out
of nine study species (Wright & Cornejo 1990).
There is mixed evidence on whether water supply
substantially limits growth in forests growing in asea-
sonal climates, where annual variation in soil water
availability is minimal. At La Selva in Costa Rica (mean
annual rainfall 3859 mm), Hazlett (1987) found that
the lowest rates of girth change for two tree species,
Carapa guianensis and Goelthalsia meiantha, occurred
during the driest period of the year, and at a well-
drained site; Breitsprecher & Bethel (1990) found that
cessation of growth for five out of eight species was also
associated with the drier part of the year. However, in
the same forest, Clark & Clark (1994) found that varia-
tion in growth rate failed to correlate with variation in
rainfall over eight years, for both adult trees and juve-
niles. In another forest in Costa Rica (mean annual
rainfall 3970 mm), a negative correlation was found be-
tween rainfall and shoot growth rate of a liana species,
Passiflora pittieri (Longino 1986).
All the studies described above concentrate on sin-
gle sites; forest growth in the tropics has rarely been
compared along rainfall gradients. However, Harring-
ton et al. (1995) examined the growth of Acacia koa
along an altitudinal gradient in Hawaii, where annual
rainfall ranged from 850–1800 mm. These authors
found that increment in aboveground woody biomass
increased as phyllode δ13C values became more nega-
tive, but was not correlated with phyllode nutrient
concentrations, suggesting that water rather than nu-
trient availability limited growth. Soil water availabili-
ty also varies at small spatial scales, differing between
topographic positions in both evergreen and semi-de-
ciduous tropical forests (Becker et al. 1988; Daws
et al. 2002; Green & Newbery 2002; Baker et al.
2002, 2003) but the influence of these differences on
tree growth has rarely been studied. However, topo-
graphic variation in soil water availability does influ-
ence plant water relations and patterns of cambial ac-
tivity in seasonal forests. Becker et al. (1988) demon-
strated that during the dry season in a semi-evergreen
forest in Panama, pre-dawn water potentials of the
shrub Psychotria horizontalis and of saplings of the
Perspectives in Plant Ecology, Evolution and Systematics (2003) 6, 21–36
tree Trichilia tuberculata, were half as low in slope
compared to plateau positions. In addition, in season-
ally dry forest in Costa Rica, species such as Astroni-
um graveolens, which occur in moist localities, are
able to maintain cambial activity into the dry season
and are leafless only for a short period, whereas girth
increment ceases during the dry season in deciduous
species in drier sites (Borchert 1999). In both exam-
ples, greater water availability in lower topographic
positions improves conditions for tree growth.
Irradiance
Irradiance declines exponentially with decreasing
height in tropical forests (Yoda 1978) and increased
canopy illumination has a clear, positive effect on tree
growth. For example, in moist evergreen forest in
Panama, saplings of ten species showed greater height
growth in high-light environments (King 1994) and at
La Selva, Costa Rica, positive correlations were found
between growth rate and crown illumination category
for eight species in size classes up to 30 cm diameter
(Clark & Clark 1992). For large trees, many studies
have used Dawkins’ (1958) classification of tree crown
position in the canopy to show effects on tree growth
(e.g. Korsgaard 1986; Silva et al. 1995; Alder & Silva
2000). For example, in individuals of Carapa
nicaraguensis, 30–40 cm in diameter in swamp forest
in Costa Rica, Webb (1999) showed that median di-
ameter increments were significantly higher in higher
crown-score classes. Also, in a study of 15 species in
tropical forest in Puerto Rico (mean annual rainfall
range 2500–4500 mm), Parresol (1995) demonstrated
that maximum growth rates were five times higher in
dominant trees than in trees in the suppressed crown-
class categories.
Variation in irradiance also influences temporal pat-
terns of growth, and may be particularly important in
aseasonal forests, where water availability is not limit-
ing. For instance, in the absence of any correlation of
growth with rainfall at La Selva, Costa Rica (mean an-
nual rainfall 3659 mm), Clark & Clark (1994) sug-
gested that inter-annual variation in irradiance deter-
mined the consistent long-term growth patterns of
adult and juvenile trees. In addition, studies of pheno-
logy have indicated that trees may time the production
of new leaves during the months of the highest irradi-
ance, if water supply is not limiting. Wright & van
Schaik (1994) found that in two weakly seasonal
forests, at La Selva, in Costa Rica and at the Ducke
Reserve in central Brazil, the number of species that
centred leaf production in the three-month period of
highest irradiance was significantly greater than ex-
pected by chance. Also, they found that in four season-
al forests, two thirds of deep-rooted species flushed

during the dry season, when irradiance was highest.
Finally, results from a biogeochemical model of forest
productivity also suggested that light may limit growth
in aseasonal forests, as a decline in modelled estimates
of net primary productivity at the wettest sites along a
transect in Brazilian Amazonia was attributed to re-
duced irradiance (Potter et al. 1998).
Nutrients
Most tropical forests grow on relatively nutrient-poor
soils. The effect of soil fertility on growth has therefore
been a focus of much experimental work, and fertilisa-
tion experiments on adult trees in natural forests pro-
vide some evidence that nutrient supply does limit
growth rates. In montane forest in Venezuela, Tanner
et al. (1992) demonstrated that trunk growth of all
species approximately doubled in plots fertilized with
N and P, and in montane forest in Colombia, N and P
fertilization also significantly increased growth for two
out of three species (Cavelier et al. 2000). On soils of
volcanic origin in montane forest in Hawaii, Vitousek
et al. (1993) demonstrated that N limited growth on
the two youngest sites (<30 and 200 years old), while P
was limiting on older soils (2000 years). However, few
studies have directly examined nutrient limitation in
lowland tropical forests. In one study in Kalimantan
(mean annual rainfall c. 3600 mm), N and P fertilisa-
tion over four years had no effect on trunk growth, but
did increase litterfall (Mirmanto et al. 1999).
The few field studies comparing patterns of soil nu-
trient availability with variation in adult tree growth
in natural forest have failed to find significant correla-
tions. Clark et al. (1998), working in forest in an asea-
sonal climate in Costa Rica with adults of nine canopy
or emergent tree species, found no difference in
growth rate between two well-drained soil-types that
differed in available P concentration by a factor of
two. In addition, Ashton & Hall (1992) found no cor-
relation between the proportional diameter growth
(1965–1985) of large trees (dbh >30 cm), and soil nu-
trient status across 13 plots in high-rainfall forests in
Borneo. However, neither of these studies compared
growth rates across the large differences in soil fertility
that are found along regional gradients of rainfall, and
differences in site fertility on this scale do influence
plantation performance. In stands of teak (Tectona
grandis) more than 10 years old in West Africa, for ex-
ample, total N in the topsoil (0–10 cm) and rooting
depth were the most important factors determining
variation in growth (Drechsel & Zech 1994). Also, for
plantations of Terminalia ivorensis across seven forest
reserves in Ghana (mean annual rainfall 1280–1650
mm), soil total N and C concentrations, cation ex-
change capacity and exchangeable Ca and Mg concen-
Variation in tropical forest growth rates 25
trations were significantly negatively correlated with
the degree of die-back, a condition that leads to wilt-
ing, chlorotic leaves, poor growth and high mortality
(Agyeman & Safo 1997). Plantation studies have also
indicated that variation in nutrient supply may be im-
portant over topographic gradients. Variation in
growth of teak over a catena in the rain forest zone of
Liberia (mean annual rainfall 2553 mm) was related
most strongly to variation in soil pH and percentage
base saturation (Zech & Drechsel 1991).
Experimental studies of seedling growth also
demonstrate that regional and topographic gradients
of soil fertility can cause differences in growth of indi-
vidual species. Veenendaal et al. (1996a) studied
seedling growth in a range of irradiances in well-wa-
tered pots of soils from semi-deciduous (pH >6.1) and
evergreen forest (pH <4.9) in Ghana, and found that
two out of 15 species grew faster on soil from semi-de-
ciduous forest, and one species grew faster on soil
from evergreen forest. The greater growth of species
on generally more fertile soils seems likely to be caused
by nutrient availability: foliar concentrations of P, N,
Ca and Mg were generally higher in seedlings grown
on soil from semi-deciduous forest (Veenendaal et al.
1996a). In contrast, the greater growth of one species
on generally less fertile soils was tentatively attributed
to a specific requirement for a trace element, such as
Cu or Zn (Veenendaal et al. 1996a). In another study,
in a comparison of growth in seedlings transplanted
into soils collected along a topographic gradient in
aseasonal forest in Sri Lanka, five out of eight species
of Shorea showed greater dry mass after two years
when grown on valley soil as opposed to ridge soil,
and seven species grew more on valley soil than on
midslope soil (Gunatilleke et al. 1996). Valley soils
had a higher pH than both mid-slope and ridge soils,
and higher concentrations of total N & Mg, and the
differences in growth were attributed to topographic
variation in soil nutrient availability.
Interactions between resources
Previously, we have considered individual resources as
if they act independently. However, variation in irradi-
ance, nutrients and water availability can have com-
plex interacting effects on tree growth. For example,
the response of plant growth to nutrient availability
increases at higher flux densities of irradiance, because
faster growth imposes an increased demand for nutri-
ents (Peace & Grubb 1982). For tropical trees, this
positive interaction between irradiance and nutrient
supply has been shown in studies of pot-grown
seedlings (Thompson et al. 1992; Lehto & Grace
1994; Veenendaal et al. 1996a; Huante et al. 1998a,b).
The demonstration of similar interactions in fertilisa-
Perspectives in Plant Ecology, Evolution and Systematics (2003) 6, 21–36
26 T. R. Baker et al.
tion experiments in the field appears to depend on the
inherent fertility of the soil and the magnitude of dif-
ference in irradiance between gap and understorey
treatments. For example, Dalling & Tanner (1995)
found a response to fertilisation of seedlings of three
species transplanted onto nutrient-poor landslides in
Jamaica under high irradiance, but no effect of fertili-
sation in the understorey. However, Fetcher et al.
(1996) found no difference in the effect of fertilisation
in a comparison of gap centre and gap edge sites on a
landslide in Puerto Rico. In contrast, Denslow et al.
(1990) found no effect of fertilisation in either gap or
understorey conditions for seven species of shrubs on
more fertile soils at La Selva, Costa Rica.
Other interactions, between light and water supply,
or water and nutrient supply have received even less
attention, even for pot-grown seedlings, and there
have been very few field-based studies. However, Fish-
er et al. (1991) showed a positive interaction between
water supply and irradiance on height growth and leaf
area production of Virola surinamensis seedlings in an
irrigation experiment in semi-evergreen forest in Pana-
ma. An interaction between nutrient supply and water
availability may be important in determining differ-
ences in the spatial pattern of tree growth observed in
different years. Baker et al. (2003) studied diameter
growth of trees > 20 cm diameter of two species over
two years in semi-deciduous and evergreen forest in
Ghana. During the first year, when there was similar,
low rainfall in both sites, growth did not differ be-
tween forests. However, during the second year, when
soil water availability was generally higher at both
sites, growth was greater in the semi-deciduous than in
the evergreen forest. This difference between the two
sites, apparent only when there was sufficient rainfall,
was attributed to the higher soil nutrient availability in
semi-deciduous forest.
In summary, within tropical rain forests, it is evi-
dent that water availability is an important influence
on tree growth rates, particularly in seasonal forests,
where it determines both the inter- and intra-annual
patterns of growth. However, it is less clear whether
the substantial spatial variation in soil water availabil-
ity between different tropical forests, or the smaller
fluctuations in soil water availability within aseasonal
forests, determine important differences in tree
growth. Variation in irradiance is the primary factor
that limits plant growth within forest stands, and pos-
sibly over long timescales for aseasonal forest as a
whole. Variation in soil fertility appears to be impor-
tant for determining variation in the growth rates of
some species, but may have a smaller role in control-
ling overall stand-level patterns of growth. However,
truly broad-scale comparisons from a large number of
sites are currently lacking.
Perspectives in Plant Ecology, Evolution and Systematics (2003) 6, 21–36
Tree growth and functional groups
We now consider variation in tree growth from the per-
spective of differences between functional groups. A
functional group can be defined as a suite of species
that share similar species-specific patterns (i.e. evolved
traits, not phenotypic) of resource use, a similar re-
sponse to disturbance, or a similar range of growth,
mortality and recruitment rates (Gitay & Noble 1997).
By recognising groups of species that share suites of co-
varying characteristics we may be able to scale-up from
the well-studied responses of few species, to model
community- or stand-level responses to perturbations
such as climate change or tree harvesting (Vanclay
1994; Condit et al. 1996). The main purpose of recog-
nising aggregations of traits in life histories is thus to
provide a semi-quantitative foundation for these prac-
tical objectives, while in practice most ecologists em-
phasise continua rather than discontinuities in the
spectrum of life histories of coexisting species.
Turner (2001) identified regeneration class and
maximum height as the primary factors that differenti-
ate tropical tree species, and below we review briefly
the evidence for trends in growth rate and response to
resource availability among functional groups defined
in these ways. Finally, we propose a new way of defin-
ing a functional group, based on differences among
species in their distribution at large spatial scales in re-
spect of soils and climate, and illustrate differences
among groups defined in this way for trees growing at
one site in semi-deciduous forest in Ghana.
Groups defined by regeneration strategy
The major axis of differentiation in ecological charac-
teristics used to separate species in tropical forests has
been related to light requirements for regeneration, be-
cause light is the primary limiting factor for growth
rate within a forest stand, and because it is possible to
allocate species on the basis of field observation alone.
Species that require the high light environment of a
gap for seed germination and/or establishment (‘pio-
neers’), are separated from species that are able to es-
tablish under canopy shade (‘non-pioneers’) (Swaine
& Whitmore 1988; Whitmore 1989). These two cate-
gories mark sections on what is now agreed to be a
continuum of responses to the range of light environ-
ments found on the forest floor (Swaine & Whitmore
1988; Alvarez-Buylla & Martinez-Ramos 1992).
Many studies have examined growth responses of
tropical tree seedlings with differing regeneration strate-
gies to variation in irradiance (e.g. Thompson et al.
1992; Lehto & Grace 1994; Veenendaal et al. 1996a).
This work has shown that under high-light conditions,
pioneers have higher growth rates than non-pioneers.

For example, in a comparison of Ghanaian tree
seedlings, three pioneer species (Terminalia ivorensis,
Milicia excelsa and Albizia zygia), all had higher relative
growth rates than a group of five non-pioneer shade
bearers (Strombosia glaucescens, Cynometra ananta,
Guarea cedrata, Celtis mildbraedii and Chrysophyllum
pruniforme) at 16% of ambient irradiance (Veenendaal
et al. 1996a). At lower light levels, whether pioneers
have higher growth rates than non-pioneers appears to
depend on exactly how the comparison is performed. A
compilation of the results of seedling growth studies of
194 species grown under contrasting light regimes
found that pioneers and light demanding non-pioneer
species had higher relative growth rates than more
shade-tolerant species, at up to 5% irradiance
(Veneklaas & Poorter 1998). However, 5% irradiance is
greater than the compensation point of all tropical trees
so far tested, and when lower irradiances are included in
experiments (e.g. Agyeman et al. 1999) pioneers are
found to grow more slowly than shade-bearers, and typ-
ically show negative growth at 2% irradiance. The
timescale of the experiment also influences the interspe-
cific differences that are observed (Sack & Grubb 2001).
When measured over short periods in low irradiance,
seedlings of light demanding species may have higher
growth rates than more shade-tolerant species, due to
the higher growth relative growth rates associated with
smaller seed sizes. However, over longer timescales,
shade-tolerant species are expected to out-perform light
demanders (Sack & Grubb 2001).
Amongst adult trees in the field, pioneer species are
usually found to grow faster than more shade-tolerant
species, presumably because they both have higher in-
trinsic growth rates at a given irradiance, and because
they are found in high-light sites. Swaine (1994) re-
ported growth rates of pioneer trees in semi-deciduous
forest in Ghana more than double those of any more
shade-tolerant group, and in semi-evergreen forest in
Panama, Condit et al. (1996) found that species ‘colo-
nizing index’, defined as the proportion of recruits
found in light gaps, was positively correlated with
species growth rates in the 10–20 cm diameter class.
However, adult tree growth rate is not always simply
related to regeneration strategy. For example, Milton
et al. (1994), working in semi-evergreen forest in
Panama, compared the growth rate of two groups of
species, defined on the basis of whether recruitment
was significantly higher, or did not differ, between low
and high canopy sites, based on previous work by
Welden et al. (1991). Over 13.6 years, for trees >19.1
cm diameter, growth rates were significantly higher in
species whose regeneration was not related to low
canopy sites. This complexity is at least partly caused
by the considerable variation in growth rate that is
found in non-pioneer species beyond the seedling stage
Variation in tropical forest growth rates 27
due to changes in species light requirements during on-
togeny (Clark & Clark 1992, 1999). For example,
many species that are shade-tolerant as seedlings re-
quire a large increase in canopy illumination as
saplings to reach the canopy (Jones 1956; Clark &
Clark 1992; Hawthorne 1995). In addition, others
have noted small-stature species whose seedling estab-
lishment requirements match those of pioneers, but
which later in life tolerate deep shade, and behave as
shade-tolerant understorey trees, such as the ‘cryptic
pioneers’ of Hawthorne (1995, p. 16). An analogous
ontogenetic shift in light requirement is displayed by
the small-seeded canopy tree Alseis blackiana on Barro
Colorado Island, Panama, which establishes only in
canopy gaps but persists for several years in the shade
after canopy closure (Dalling et al. 2001).
In addition to higher growth rates at high irradi-
ance, pioneer species also have greater photosynthetic
plasticity and show greater growth responses to in-
creased irradiance than shade-tolerant species. A com-
parison of the range of maximum photosynthetic
rates, under high and low light, for species subjectively
described as early- and late-successional (16 and 24
species, respectively), showed that early-successional
species have a greater range in Amax values than late-
successional species, because of significantly higher
values under high light (Strauss-Debenedetti & Bazzaz
1996; Thomas & Bazzaz 1999). For adult trees in the
field, Welden et al. (1991) working in a semi-evergreen
forest in Panama, noted that increased growth in low
compared to high canopy sites was a particular feature
of six pioneer species.
Nutrient addition experiments with seedlings show
similar patterns to those with irradiance, with a
greater response in pioneer and early-successional
species. Huante et al. (1995) demonstrated that three
early-successional species from a deciduous forest in
Mexico showed a greater proportional increase in
biomass (3.7–124 fold increase) between low and high
(0 and 41 ppm) P treatments, than four late-succes-
sional species (1.2–2.4 fold increase). In addition,
Raaimakers & Lambers (1996) showed that the
biomass of a pioneer species from Guyana, Tapirira
obtusa doubled over six months at high (>250 mg P
per plant) compared to low (<10 mg P per plant)
P-supply, whereas growth of the non-pioneer species
Lecythis corrugata was not affected. In natural forest
on a landslide in Puerto Rico, Fetcher et al. (1996)
found that two pioneer species showed a greater re-
sponse in growth to N and P fertilisation than two
shade-tolerant species. Also, in lowland forest in Puer-
to Rico, saplings of the pioneer Cecropia schreberiana
were three to five times more numerous in fertilized
compared to control plots after four years (Walker
et al. 1996).
Perspectives in Plant Ecology, Evolution and Systematics (2003) 6, 21–36
28 T. R. Baker et al.
Groups based on maximum height
A second axis of differentiation amongst tree species in
tropical rain forest, related to the vertical gradient in
irradiance, is caused by differences in maximum
height. Species with low maximum heights typically
have low growth rates. For example, in semi-decidu-
ous forest in Ghana, Swaine (1994) found that mean
annual diameter-increment of large shade-tolerant
species was double that of small-stature species, and in
Sungei Menyala Forest Reserve in Malaysia,
Manokaran & Kochummen (1987) found that mean
annual diameter increments of understorey species
were less than 2 mm yr–1, compared to increments
greater than 3 mm yr–1 for emergent species. Growth
rates are also less variable in species with low maxi-
mum heights. At La Selva, Costa Rica, Lieberman &
Lieberman (1987) demonstrated that total variation in
growth rate is approximately five times greater in
shade-tolerant canopy species than in understorey
species.
Distinguishing whether these patterns are caused by
a correlation between tree diameter and absolute mea-
sures of diameter growth, lower levels of crown illumi-
nation for smaller trees, or because small trees have
lower inherent growth rates under similar levels of ir-
radiance, requires comparisons that control for tree di-
ameter and the light environment. In Pasoh Forest Re-
serve, Malaysia, using asymptotic height derived from
height/diameter relationships as an estimator of maxi-
mum species height, Thomas (1996) found a linear
correlation (r2 = 0.56, P < 0.001) between maximum
height and mean annual growth rate of adult trees for
38 species. In addition, the same positive correlation
(r2 = 0.20, P < 0.01) was found using the growth rates
of saplings (1–2 cm diameter) rather than adult trees,
suggesting that the relationship is not confounded by
variation in tree diameter. Also, in a comparison of
photosynthetic characteristics of similar-sized saplings,
under similar light conditions, of species with different
asymptotic heights, smaller-statured species were
found to have an inherently lower leaf level photosyn-
thetic capacity (Amax), compared to species with
greater maximum height, within the same genus
(Thomas & Bazzaz 1999).
Both inherently lower diameter growth rates and
lower levels of crown illumination therefore charac-
terise species with low maximum heights. However, it
is important to note that all of the above studies refer
predominately to shade-tolerant tree species. These
species establish in the understorey and therefore ex-
perience increasing light levels as they increase in size.
This pattern will inevitably lead to increased growth
rates in larger individuals, and, on an evolutionary
timescale, favour increased capacity for growth in tall-
Perspectives in Plant Ecology, Evolution and Systematics (2003) 6, 21–36
statured species (Thomas & Bazzaz 1999). In contrast,
pioneer species establish under high light conditions,
so there will not be a positive correlation between light
availability and plant size. Therefore, similar ecologi-
cal or evolutionary arguments for faster growth in
species with greater maximum height are unlikely to
apply to pioneers (Thomas & Bazzaz 1999).
Groups defined by species’ associations
with specific environmental conditions:
a test from the forest zone of Ghana
A third axis of differentiation of tropical forest species
may relate to the associations between species’ distri-
butions and climatic and edaphic factors. Species’ dis-
tribution in respect of edaphic factors has been
demonstrated to be a useful predictor of growth rates
in studies of temperate plants, where species from dif-
ferent habitats have been grown under the same envi-
ronmental conditions. The generalisation to emerge
has been that species from nutrient-poor habitats typi-
cally have lower maximum growth rates than species
from nutrient-rich sites when they are grown under the
same experimental conditions (Grime & Hunt 1975;
Grime 1979; Chapin 1980; Chapin et al. 1986), al-
though this is not universally applicable (Grubb
1998). Tropical forest tree species have often been
shown to have distinctive distributions across large-
scale edaphic and climatic gradients (e.g. Gartlan et al.
1986; Newbery et al. 1986; Baillie et al. 1987;
Tuomisto et al. 1995; Swaine 1996). However, the rel-
evance of these associations to variation in growth
rate within one site has not previously been examined.
Here, we test this idea using the extensive ecological
knowledge of Ghanaian forest species, and inventory
data from a semi-deciduous forest.
Firstly, we developed a classification of tree species
incorporating both their regeneration requirements
and their association with different parts of a major
resource gradient. Within Ghanaian tropical forests,
regional scale patterns of species distribution have
been comprehensively described across a gradient of
increasing rainfall and decreasing soil fertility towards
the south-west of the forest zone (1200–2300 mm yr–1,
soil pH 7.0–3.8; Hall & Swaine 1976, 1981). The first
axis of a multivariate ordination of compositional
data from 155 closed-canopy forest plots was shown
to quantify species position along this gradient, with
decreasing axis one scores in the wettest, least fertile
forest sites (Hall & Swaine 1981). To categorise
species’ environmental preferences, the median score
(25) of all species from 23, 1-ha plots in a range of
Ghanaian forest types was used as an arbitrary thresh-
old value (Baker 2000). Species with axis one scores
above this value were classified as dry forest special-

Fig. 1. Distribution of diameter increments within Ghanaian semi-decidu-
ous forest, for six functional groups, defined by regeneration strategy and
distribution pattern with respect to climatic and edaphic conditions.
(a) Shade-bearers, (b) non-pioneer light-demanders, and (c) pioneers. There
is a significant difference in median growth rate between dry- and wet-forest
pioneer species. Box size represents the range between the first and third
quartiles, and whiskers extend 1.5 times this range. Outliers are shown as
asterisks; mean (open circle) and median (shaded circle) are also indicated.
Variation in tropical forest growth rates 29
ists, and species with scores below this value were clas-
sified as wet forest specialists. This categorization rep-
resents a coarse division of species responses to a re-
gional scale environmental gradient, and does not con-
sider local (e.g. topographic) influences of water and
nutrient availability. However, the environmental fac-
tors that determine species’ distributions at different
scales are likely to be similar. For example, in Ghana,
the abundance of Celtis mildbraedii declines as the
landscape becomes wetter, both at local and regional
scales (Swaine & Hall 1986; Hawthorne 1995).
Therefore, we are confident that our classification ade-
quately, if crudely, represents species’ environmental
preferences.
Species were also grouped according to their regen-
eration requirements (‘pioneer’, ‘non-pioneer light-de-
mander’ and ‘non-pioneer shade-bearer’), following
Hawthorne (1995). Therefore, overall, species were
grouped into six categories, each including species
with a range of maximum heights.
Inventory data were obtained from permanent sam-
ple plots located in Bobiri Forest Reserve, in Ghana
(1°15–23′W, 6°40–42′N), which is classified as Moist
Semi-deciduous Forest (Hall & Swaine 1981). Mean
annual rainfall (1961–1993) for Kumasi, 15 km west
of the forest is 1183 mm. The main dry season occurs
from December to March (Swaine et al. 1997), with a
less severe drier period from July to September (Vee-
nendaal et al. 1996b). Most of the reserve contains
soils developed over upper and lower Birrimian phyl-
lite (Adu 1974), with gently undulating topography.
The soils consist of red, silty clay loams in summit
areas and become paler in colour lower downslope.
Valley soils are grey, sandy loams and clays (Adu
1974).
Seven 1-ha plots that had not been damaged by fire
or degraded by logging were selected in a range of to-
pographic positions. They were established by the
Ghana Forestry Department in July/August 1990 and
re-enumerated in July/August 1996. The girth of each
tree >20 cm dbh was measured to the nearest mm and
each measured tree was scored for the degree of crown
illumination (‘crown score’) on a five point scale
(Dawkins 1958). The painted point of measurement
was at a height of 1.3 m, or 50 cm above the top of
any buttress. Annual diameter growth of surviving
trees was compared between wet- and dry- forest
species for each of the three regeneration categories
using Kruskal-Wallis tests.
For the three comparisons, the only significant dif-
ference in growth rate was found between wet- and
dry-forest pioneer species (Fig. 1). Dry-forest pioneer
species had significantly higher median growth rates
than wet-forest pioneers (H = 6.25, P < 0.012, n = 92,
dry-forest pioneers, and 27, wet-forest pioneers). The
Perspectives in Plant Ecology, Evolution and Systematics (2003) 6, 21–36
30 T. R. Baker et al.
Fig. 2. Annual diameter increment as a function of tree diameter for wet and dry forest pioneers, within semi-deciduous forest in Ghana.
difference in growth rate between these groups does
not appear to be caused by differences in tree size or
crown illumination. Wet- and dry-forest pioneers com-
prise trees of a similar diameter range, and there is no
significant difference in the median diameter between
the two groups (Fig. 2; H = 0.52, P > 0.05). In addi-
tion, the proportion of trees with high levels of crown
illumination are similar: 30% of dry forest pioneers
and 29% of wet forest pioneers were classified as
Dawkins’ crown score 4 or 5.
The lower growth rates of wet-forest compared to
dry-forest pioneers may reflect an inability of the wet
forest species, that typically occur on less fertile soils,
to respond to the higher nutrient status of the soils of
the semi-deciduous forest site. This contrast is sup-
ported by seedling experiments on two pioneer species
that have different distribution patterns, Triplochiton
scleroxylon and Lophira alata (Table 2). Both species
are strongly light demanding, and attain large sizes,
greater than 90 cm dbh (Hawthorne 1995). However,
they show strongly contrasting patterns of seedling
growth on low and high fertility soils. Under well-wa-
tered conditions, T. scleroxylon showed significantly
higher growth on soil from semi-deciduous compared
to evergreen forest sites, whereas L. alata grew fastest
on the less fertile, evergreen forest soil (Swaine et al.
1997; Veenendaal et al. 1996a). With the results here,
these patterns suggest that there may be variation in
the maximum growth rate of pioneer species in
Ghanaian forest, related to their association with par-
ticular edaphic and climatic conditions.
In summary, in terms of growth rate, the classifica-
tions of functional groups proposed by Turner (2001)
differentiates groups of similar species. Classifications
Perspectives in Plant Ecology, Evolution and Systematics (2003) 6, 21–36
based both on regeneration strategy and maximum
size, generally divide species that have low growth
rates and show little response to increasing resources,
from more responsive, faster-growing species. Howev-
er, in both cases, exceptions occur. The growth rate of
adult trees of non-pioneer species may be modified by
variation in light demand during ontogeny, and maxi-
mum size may not be associated with any systematic
variation in growth rate for pioneer species. Classifica-
tions of species based on their association with partic-
ular edaphic and climatic conditions may be useful for
categorizing important variation in the ecology of pio-
neer species in tropical forests.
Functional gradients in forest composition
So far, we have focussed on two rather different per-
spectives on tropical tree growth, and separately con-
sidered variation in growth rates with resource avail-
ability, and between functional groups. However, both
factors will be important for understanding spatial
variation in stand-level growth rates, if there are dif-
ferences in the relative abundance of functional groups
between forests. There is increasing evidence of such
functional gradients at a range of scales within tropi-
cal forests.
Studies of gradients in the functional composition
of tropical forests have concentrated on variation in
the relative abundance of pioneer and non-pioneers, or
variation in stand-level mean values for traits such as
wood density, that are strongly correlated with
species’ light demand (Whitmore 1998). For example,
in Ghana, pioneer species are both most abundant and
 Variation in tropical forest growth rates 31

Table 2. Dry- and wet-forest pioneers in Ghanaian tropical forest. Nomenclature and classification of pioneers follows
Hawthorne (1996); association with dry or wet forest based on a multivariate ordination of species presence or absence in
155 plots across the forest zone of Ghana (Hall & Swaine 1981).

Dry forest pioneers Wet forest pioneers

––––––––––––––––––––––––––––––––––––––––––––––––– ––––––––––––––––––––––––––––––––––––––––––––
Species Family Species Family

Baphia pubescens Fabaceae Cola caricifolia Sterculiaceae

Canarium schweinfurthii Burseraceae Daniellia ogea Fabaceae
Ceiba pentandra Bombacaceae Daniellia thurifera Fabaceae
Celtis adolfi-friderici Ulmaceae Hannoa klaineana Simaroubaceae
Cleistopholis patens Annonaceae Lophira alata Ochnaceae
Draceana arborea Agavaceae Petersianthus macrocarpus Lecythidaceae
Ficus exasperata Moraceae Bombax brevicuspe Bombacaceae
Holoptelea grandis Ulmaceae Zanthoxylum gillettii Rutaceae
Lannea welwitschii Anacardiaceae
Milicia excelsa Moraceae
Morus mesozygia Moraceae
Newbouldia laevis Bignoniaceae
Rauvolfia vomitoria Apocynaceae
Bombax buonopozense Bombacaceae
Ricinodendron heudelotii Euphorbiaceae
Stereospermum acuminatissimum Bignoniaceae
Terminalia superba Combretaceae
Tetrapleura tetraptera Fabaceae
Tetrorchidium didymostemon Euphorbiaceae
Triplochiton scleroxylon Sterculiaceae
Zanthoxylum leprieurii Rutaceae

diverse in semi-deciduous forests, where rainfall is
lower and soils are more fertile than in evergreen for-
est (Hawthorne 1996). In addition, mean stand level
wood density and seed size are lowest in southern
compared to central Guyana, indicating that southern
forests have a greater relative abundance and diversity
of more light-demanding species (ter Steege & Ham-
mond 2001). Also, in a comparison of 59 plots across
Amazonia, mean stand level wood density was 12%
higher in eastern, compared to north-western Amazon
forests (Baker et al., in press).
Variation in past and current rates of disturbance
are probably the most important factor determining
the variation in the composition of tropical forests
with respect to light demand. For both Ghana and
Guyana, higher proportions of light-demanding taxa
are associated with areas that have a history of higher
rates of anthropogenic disturbance (Fairhead & Leach
1998; ter Steege & Hammond 2001). Past human ac-
tivity has also been suggested as an important factor
determining the pioneer-rich flora of Barro Colorado
Island in Panama (Sheil & Burslem 2003). Across
Amazonia, it seems unlikely that there are important
systematic differences in past human activity that have
generated the observed variation in stand-level wood
density (Baker et al., in press). Here, higher abiotic dis-
turbance rates in western compared to eastern Ama-
zon forests may explain the pattern. However, it is also
likely that soil nutrient availability influences the func-
tional composition of tropical forest, as the growth
rates of pioneer species are particularly responsive to
higher soil nutrient concentrations. The forests of both
Ghana and Guyana that have a higher abundance of
light demanding taxa, occur on more fertile soils
(Hawthorne 1996; ter Steege & Hammond 2001). As
this pattern coincides with variation in the intensity of
past human disturbance, it is difficult to distinguish
the relative importance of the two factors.
These functional gradients in tropical forest compo-
sition have important implications for spatial varia-
tion in forest growth and dynamics, as light demand is
generally positively correlated with species’ growth
and mortality rates (Whitmore 1998). For example, at
a small scale, Tim Whitmore’s long-term study of for-
est dynamics on Kolombangara in the Solomon Is-
lands has yielded evidence of spatial variation in
stand-level turnover rates that correlates with varia-
tion in functional group composition (Burslem &
Whitmore 1999, in press). In addition, within a net-
work of plots in aseasonal climates across Sarawak, a
guild of pioneer species increased in abundance at a
high-nutrient site, but were rare elsewhere, resulting in
positive correlations between soil nutrient concentra-
tions and diameter growth of recruits (Ashton & Hall
1992). At a larger scale, the lower wood density of
western compared to eastern Amazon forest is associ-

Perspectives in Plant Ecology, Evolution and Systematics (2003) 6, 21–36

32 T. R. Baker et al.
ated with higher tree turnover rates (Phillips et al.
1994, in press) and higher stand-level wood produc-
tion, calculated on both a volume and mass basis (Y.
Malhi et al., unpubl. results). Finally, in a transconti-
nental comparison of the dynamics of an evergreen
and a semi-deciduous forest, a guild of pioneer species
present only in the semi-deciduous forest was suggest-
ed to determine overall plot differences in growth and
mortality (Condit et al. 1999). The more seasonal for-
est had higher overall growth and mortality rates than
a plot lacking the guild of fast-growing pioneer
species.
In summary, these studies demonstrate that spatial
variation in the proportion of light-demanding taxa,
occurs at range of scales in tropical forests. Variation
in disturbance rates appears to be the main factor that
determines these patterns, although soil nutrient avail-
ability may also be important. The combined effects of
functional group composition and resource availabili-
ty will therefore determine spatial variation in stand-
level growth.
Future directions
In this review, we have used a broad definition of
‘growth rate’ incorporating both changes in size and
carbon content of plants, plant parts and populations.
Size is important, both ecologically as an important
determinant of competitive success, and for silvicul-
ture as timber yields are typically determined on the
basis of volume. In contrast, growth rates calculated in
terms of changes in carbon content are more closely
related to variation in rates of photosynthesis and res-
piration and hence resource availability, and are im-
portant for calculating stand-level carbon balance.
The two quantities are related through variation in al-
location strategies, which vary both along resource
gradients and between functional groups. In terms of
aboveground growth, inclusion of wood density in cal-
culations of the growth rate of adult trees is essential
for understanding how different life history strategies
compare in terms of carbon uptake (e.g. Enquist et al.
1999). In addition, more work is required on how
trends in belowground allocation correspond to varia-
tion in aboveground productivity. For example, fine-
root densities in the topsoil are higher in tropical
forests on highly infertile soils (Coomes & Grubb
2000), but across Amazonia it is not known to what
extent these kind of belowground patterns may offset
spatial variation in aboveground productivity (Y.
Malhi et al., unpubl. results).
Comparative studies of the ecology and dynamics
of a large number of forests are required to understand
the combined effects of functional groups and resource
Perspectives in Plant Ecology, Evolution and Systematics (2003) 6, 21–36
availability on spatial variation in forest growth rates.
These studies will need to account for variation in
water and nutrient availability as well as variation in
light. In addition, quantifying the variation in life-his-
tory strategies using appropriate traits, such as wood
density and maximum size, will allow tests of whether
effects of resource availability are independent of vari-
ation in species composition. The ideas of Tim Whit-
more in defining and exploring functional differences
between tropical forest trees will therefore remain cen-
tral to future research into this fundamental aspect of
tropical rain forest ecology.
Acknowledgements. For advice and assistance during field-
work in Ghana we thank Dr. V.K. Agyeman (FORIG, Ku-
masi) and Dr. T.K. Orgle (Forest Management Support Cen-
tre, Kumasi). We also thank Kofi Affum-Baffoe, Yaw Atua-
hene and Raymond Votere (Inventory Unit, Forest Manage-
ment Support Centre, Kumasi) for compiling the forest in-
ventory data used in this study. Peter Grubb, Ian Turner and
Oliver Phillips provided very helpful comments on a previ-
ous version of this manuscript. This work was largely fund-
ed by a University of Aberdeen Faculty studentship to TRB,
who also acknowledges current financial support from the
Max-Planck-Institut für Biogeochemie, Jena, Germany.
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