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Beyond the regeneration phase:

differentiation of height-light
trajectories among tropical tree
species
Lourens Poorter
Journal of Ecology

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Journal of
Beyond the regeneration phase: differentiation of height–
Blackwell Publishing, Ltd.

Ecology 2005
93, 256–267 light trajectories among tropical tree species
LOURENS POORTER, FRANS BONGERS, FRANK J. STERCK and
HANNSJÖRG WÖLL *
Forest Ecology and Forest Management Group, Wageningen University, PO Box 47, 6700 AA Wageningen, the
Netherlands, and *Sommersbergseestr. 291, A-8990 Bad Aussee, Austria

Summary
1 A height–light trajectory (HLT, a fitted curve relating canopy exposure to tree height)
was determined for populations of individuals of each of 47 tree species in a Liberian
lowland rainforest. The HLTs were compared and related to tree allometry and adult
stature. Crown exposure was measured for 7460 trees and related to tree height using a
multinomial regression analysis. Individual trees were followed for 2.8–9.8 years.
2 The trajectories of the 47 species were compared with the average vertical light profile
in the forest canopy. Evidence was found for the existence of all nine trajectories hypoth-
esized on the basis of three possible light environments (high, intermediate and low) for
juveniles and adults. The classical paradigm of pioneer vs. shade tolerant, based on seed
and seedling responses, does not therefore apply to post-seedling stages.
3 The majority of the species followed the vertical light profile in the forest canopy,
starting in low light environments in the juvenile stage and ending up in high light
environments in the adult stage. Only two species complied with the classic notion of
whole-life shade tolerants and whole-life shade intolerants (one each).
4 The predictable vertical light gradient in the forest canopy has led to a close associ-
ation between adult height, light trajectories and allometric traits. Large-stature species
tend to have relatively slender stems and narrow crowns, and therefore realize a faster
temporal and height-related increase in crown exposure.
5 Tree species have different height–light trajectories when they grow from seedling to
adulthood. This may have profound repercussions for our current views on plasticity and
adaptation, light partitioning and species coexistence, and on silviculture and management.
Key-words: architecture, functional groups, Liberia, light, niche partitioning, regener-
ation, tree, tropical rain forest, shade tolerance
Journal of Ecology (2005) 93, 256–267
doi: 10.1111/j.1365-2745.2004.00956.x

seedlings of tree species are found. However, species

Introduction
Forest succession, composition and dynamics are often
explained based on the light requirements of tree
species for establishment, survival and growth (Finegan
1984; Pacala et al. 1996). Traditionally, foresters in both
temperate and tropical zones have classified tree spe-
cies into groups according to their shade tolerance in
the regeneration phase, leading to the classical dichot-
omy of pioneer vs. shade-tolerant species (Swaine &
Whitmore 1988). Strong early environmental require-
ments in combination with high mortality rates may
determine when, where and under what conditions
© 2005 British Correspondence: Lourens Poorter
Ecological Society (e-mail lourens.poorter@wur.nl).
may differ in their tolerance of shade at successive stages
in their life cycle (Oldeman & van Dijk 1991; Grubb
1996; Bazzaz 1998). The ‘regeneration niche’ of a spe-
cies, as defined by Grubb (1977), embraces all stages in
the regeneration process, and up till now there have
been very few quantitative studies of the requirements
of tree species beyond the juvenile stage (e.g. Clark &
Clark 1992).
While every tree may have its own unique light tra-
jectory from seed to adult tree, a general classification
can be used to indicate the possible alternative pathways
during a tree’s life (individual ontogenetic trajectories).
In Fig. 1 we simplify these possibilities by consider-
ing four critical life-history stages and three light
levels. After dispersal, a seed may be found under low,
257
Height–light
trajectories of
tropical tree species
Fig. 1 Potential height–light trajectories for species in
different stages of their life cycle. Average irradiance at the
population level can be low, intermediate or high and light
levels may either remain constant from seed to adult stage
(vertical arrows) or shift from one stage to the next (diagonal
arrows). The nine possible pathways between juvenile and
adult (i.e. above the broken line) are the focus of this paper.
intermediate or high light conditions. The seedling
growing out of this seed may be found in lower, similar
or higher light conditions than the seed, as may be the
juvenile and adult stage of that individual, depending
on the changes in the individual tree’s light environ-
ment. Trajectories can be described in the same way at
the population level (population or species height–
light trajectory, HLT), where the light levels experi-
enced by seeds, seedlings, juveniles and adults may differ,
depending on the light requirements for germination
and establishment, and on the light dependency of sur-
vival, growth and reproduction. Species thus may
show different shifts in their light requirements in
different stages of their life cycle. The most extreme tra-
jectories are formed by the outer pathways in Fig. 1:
on the left, vertical arrows represent species that grow
up and mature in the low light conditions of the forest
understorey (traditionally referred to as strict shade-
tolerant species), whereas species that establish and
grow up in the bright light environment of gaps (light
demanding or pioneer species) are on the far right. In
between there is a whole gamut of potential light tra-
jectories: 81 hypothetical pathways result from only
four discrete life-history stages and three light levels
(Fig. 1) and, in reality, the two axes are continuous,
leading to an infinite number of possible pathways.
In tropical rain forest there is a strong and predict-
able vertical gradient in light availability: a progressive
exponential extinction of light occurs when incoming
radiation from above is intercepted by successive
leaf layers in the forest canopy. Trees therefore should
© 2005 British
encounter brighter light conditions as they increase in
Ecological Society, height. The null hypothesis for a height–light trajectory
Journal of Ecology, thus would be to change from shade when small to
93, 256–267 bright light when tall. What processes lead to possible
deviations of species HLTs from this null model? Spe-
cies may either deviate from the average light environ-
ment right from the beginning of their life cycle, or such
differences may only become apparent in later stages.
Experimental evidence shows that both pioneer and
non-pioneer species may germinate under a wide range
of environmental conditions (e.g. Kyereh et al. 1999;
Peña-Claros 2001; Pearson et al. 2002), contradicting
the classical ideas (cf. Swaine & Whitmore 1988). Selec-
tion occurs therefore mainly in later stages of the life
cycle, when the species are growing towards the canopy.
From a physiological point of view, trees need brighter
light conditions when growing in size: the ratio between
photosynthesizing and respiring tissue declines with
plant size, leading to an enhanced whole-plant light
compensation point (Givnish 1988). The height-dependent
changes in the whole-plant light compensation point
may differ amongst species, and the species may follow
therefore different height–light trajectories.
It is likely that the shape of the light trajectory is
related to the maximum adult stature of the species.
Many large-stature species only become reproductive
when they attain a position in the canopy and get access
to light (e.g. Thomas 1996a; Zuidema & Boot 2002). As
a consequence, these species should grow faster towards
the canopy, and realize a faster temporal and height-
related increase in crown exposure than small-stature
species that do not have to grow to the canopy (Kohyama
& Hotta 1990). Tree allometry might play a decisive
role in this respect, as slender stems and narrow crowns
allow for a rapid vertical tree extension (Kohyama &
Hotta 1990; Sterck & Bongers 1998; Poorter & Werger
1999; Sterck et al. 2001; Kohyama et al. 2003; Poorter
et al. 2003).
Despite its central role as a working model in forest
ecology, little quantitative information is available on
the shade tolerance of tree species. Species are often
subjectively classified in two or three groups, based on
the ecological knowledge and best-educated guesses
of the researchers. The last decade has seen a growing
body of publications, in which the light environments
of tropical tree species have been quantified using
PAR sensors, hemispherical photographs, or qualitative
estimates of the light environment (e.g. Welden et al.
1991; Clark & Clark 1992; Clark et al. 1993; Lieberman
et al. 1995; Davies et al. 1998; Rose 2000; Poorter &
Arets 2003). Most studies, however, focus only on one
life-history stage, or consider only few species (but see
Hawthorne 1995). A community-wide approach is needed
to provide the necessary resolution for revealing
ecologically and statistically sound patterns, and to gain
insight into the extent to which tree species partition
the light gradient. Lieberman et al. (1995) took such a
community-wide perspective, and considered trees
over a large size range. They found that 14% of the spe-
cies indeed occurred in darker or brighter conditions
than expected, whereas 86% had a random distribution
with respect to light. However, by the nature of their
analysis, they excluded a priori the possibility that
258 species may switch light requirements with tree height
L. Poorter et al. (cf. Fig. 1).
If such height-related shifts exist, they may have pro-
found repercussions for our current views on plasticity
and adaptation (Reich 2000), light partitioning and
species coexistence (Hubbell et al. 1999), and silvicul-
ture and management (Lamprecht 1990). Species that
experience large changes in light requirements with
height might be expected to have greater trait plasticity
than species that consistently occur under high or low
light conditions (Popma et al. 1992; Grubb 1998; Reich
2000). Similarly, the influence of light partitioning
might seem weak when only the seedling stage is con-
sidered, but might be stronger when crossovers in light
requirements occur as species increase in size (Sack &
Grubb 2001). Finally, silvicultural interventions such
as liberation thinning might be more refined if we know
for what species it is needed, and at what ontogenetic
stage it should be applied.
Here we draw on a large data set from a Liberian rain
forest, in which height and crown exposure were measured
for 7460 trees. Temporal changes in crown exposure
were followed for a 6-year period. We describe the height–
light trajectories of 47 species using multinomial
logistic regression analysis. Monitored trees had a
diameter at breast height of 10 cm or over, and our ana-
lysis is therefore confined to the juvenile and adult stage
(Fig. 1).
We address two questions. First, do species differ in
their height–light trajectories from the average vertical
light profile in the forest, and if so, how common is each
of the hypothetical pathways identified? Secondly, are
differences in these trajectories related to the maximal
adult stature and allometry of the species?
Materials and methods
   
The field study was carried out in the lowland wet ever-
green forest (cf. Bongers et al. 2004) of Grebo National
Forest in south-east Liberia (5°30′ N, 7°30′ W). Total
annual rainfall is about 2000 mm, with a dry season
(< 100 mm rainfall month−1) from December to Feb-
ruary. Average canopy height is 35 m, the average stem
density is 385 stems ha −1, and species richness is 58
species ha−1 (for trees > 10 cm d.b.h.) (Poker 1993).
Twenty 1-ha permanent sample plots (PSPs) were
randomly established in an area of 900 ha, between 1978
and 1985. The nearest neighbour distance amongst
plots averages 440 m (range 210– 990 m). Fifteen to
20 years before plot establishment, selective logging
(one tree ha−1) occurred in six plots. The other plots
show no signs of recent human disturbance (Poker
1993). All trees with a diameter at breast height (d.b.h.)
© 2005 British
larger than 10 cm were measured for their diameter,
Ecological Society, tagged and identified. The height to the top of each tree
Journal of Ecology was measured with a telescope pole for heights up to
93, 256–267 15 m, and with a clinometer for heights > 15 m. Crown
width was determined as the average diameter of the
east–west and north–south cross-sections of the crown.
The crown exposure was classified on a five-point scale
(Dawkins & Field 1978): 1 = no direct light, 2 = lateral
light, 3 = partial overhead light, 4 = more than 90% of
the crown area receives direct overhead light, 5 = emer-
gent crown with direct light from all directions. Esti-
mates of this crown exposure index (CE) are repeatable
and accurate (Clark & Clark 1992) and there is a good
correlation between CE and canopy openness (Davies
et al. 1998) or the amount of incident radiation as esti-
mated with hemispherical photographs (Clark et al.
1993). The survival, diameter and CE of the trees were
re-measured after 2.8–9.8 years (mean 6.5 years).
 
The height–light trajectories were analysed with a
multinomial logistic regression (cf. Sheil, Chave, Salim,
Vanclay and Hawthorne, personal communication).
Such a regression classifies subjects based on values of
a set of predictor variables (Norusis 1997) and is more
general than logistic regression, because the dependent
variable can include more than two categories. This
analysis allows us to predict CE at first census as a func-
tion of tree height. The probability of a tree (pi) being in
a certain CE class i (i = 1, 2, 3, 4) is a function of its
height fi (h):
pi = exp( fi )/(1 + exp( f1) + exp( f2) + exp( f3) + exp( f4))
where i is 1–4.
The probability of CE class 5 can be calculated as:
p5 = 1 − p1 − p2 − p3 − p4
Several functions can be used to relate CE to the height
of the tree. We used both a simple polynomial function
(fi (h) = ai + bi × height), which allows for a monotonic
increase of CE with height, and a quadratic polynomial
function ( fi (h) = ai + bi × height + ci × height2 ), which
allows for local minima and maxima. Only the 47
species that had a sufficient number of individuals
(n ≥ 30) were included. The quadratic polynomial
function gave a significantly better fit than the simple
polynomial function for five out of 47 species. The
goodness of fit is indicated by the Nagelkerke r2, which
indicates what proportion of the variation is explained
by the regression model (Nagelkerke 1991). The pre-
dicted mean CE at any given height was subsequently
calculated as the weighted average of the five CE
classes:
CE = p1 × 1 + p2 × 2 + p3 × 3 + p4 × 4 + p5 × 5
For each species we calculated this predicted mean CE
for 5-m height intervals between 5 and 50 m height.
The 20 plots differ in their average crown exposure
(one-way , F = 36, P < 0.001). A difference in
259 HLT among species could arise if species were to occur
Height–light only in one or a few plots with extreme light levels. We
trajectories of used an index of dispersion to evaluate whether species
tropical tree species had a clumped distribution with respect to plots. For
each species the index of dispersion was calculated as
the ratio of the variance to the mean of the stem density
in the different plots (cf. Fowler et al. 1998). Most of
the species had a random or regular distribution over
the plots, but five had a significantly clumped distribu-
tion (χ2 > 33.5, P < 0.05). Of these, Stachyothyrsus
stapfiana occurred only in one plot, and Calpocalyx
aubrevillei occurred in six plots and the HLT is therefore
potentially biased. The other three species (Craterisp-
ermum caudatum, Diospyros sanza-minika and Gilber-
tiodendron preussii) occur in more than 10 plots and
plot-specific effects on HLTs are likely to average out.
Species differ substantially in the speed with which
they gain access to light by realizing an increase in
height. The height-dependent increase in CE was cal-
culated as the slope of the predicted CE against height
(in units CE per metre). We calculated the temporal
change in CE (in units CE per year) as the difference in
CE between second and first measurement, divided by
the monitoring period. Differences in temporal change
in CE among species were tested with an .
Average crown exposure and height-dependent and
temporal changes thereof were related to the maximal
tree height (Hmax) and to stem slenderness and crown
diameter of the species. The Hmax was calculated as the
95th percentile of the height values in the species’ popu-
lation, thus correcting for outliers. This is important,
because it is easy to overestimate the height of tall trees.
Stem slenderness and crown diameter were calculated,
respectively, as the tree height and crown diameter
attained by an individual with a d.b.h. of 15 cm (Poorter
et al. 2003).
An average vertical light profile of the forest was
described using the crown exposure of all 7460 trees
that were encountered in the PSPs. A 95% confidence
envelope of this light profile was created for height class
intervals of 5 m (2.5– 7.5 m, 7.5 –12.5 m, etc.). For each
height class interval the crown exposures of 17 trees
(the average number of trees per species per height class
interval) were randomly drawn from the data set and
averaged. This procedure was repeated 100 times. For
each class interval we determined the 95% confidence
interval (based on the 2.5 and 97.5 percentiles) for the
average crown exposure of trees in that class interval.
We evaluated whether the HLTs of the species dif-
fered from the average vertical light profile in the forest,
by comparing the CE at the juvenile and adult stage
with the corresponding confidence intervals. Average
crown exposure of a 5 m tall tree was used to represent
juvenile light requirements. For eight species (Table 1)
the smallest individuals were between 7.5 and 14 m tall.
© 2005 British
For these species the juvenile crown exposure was
Ecological Society, extrapolated, based on the regression equations. As
Journal of Ecology, adult light requirements we took the average crown
93, 256–267 exposure of trees that had attained their maximal
height. For each life stage we compared whether the
species had a crown exposure that was above, within or
below the 95% confidence interval for trees of the same
size class in the average profile.
All statistical analyses were carried out using SPSS 10.
Results
    , 
  
For the whole forest stand, the crown exposure increases
in a sigmoid fashion with the height of the trees (Fig. 2a).
Trees that are 5 m tall have an average crown exposure
of 1.4, and 50-m trees an average of 4. On average, trees
reach the canopy (i.e. they have a crown exposure of 3)
when they are 30 m tall. The largest increases in crown
exposure occur between 20 and 40 m height.
Individual canopy species, such as Gilbertiodendron
preussii, may follow a similar type of trajectory with a
sigmoid increase in crown exposure (Fig. 2b), but under-
storey species, such as Cola buntingii, remain mostly in
low light conditions and show a slow increase in crown
exposure (Fig. 2c). Although there are clear average
height–light trajectories, there is considerable scatter:
thus many species can be found at all possible light levels,
and species may occur in a wide range of light environments
at any given height (as shown for G. preussii in Fig. 2b).
   
We assigned the species to different light strategies,
based on their average crown exposure as juveniles and
adults, and on the switches they make during this tran-
sition between low, intermediate and high light com-
pared with the average light profile in the forest canopy
(Figs 3 and 4). Each of the nine possible light trajector-
ies proposed in the upper part of Fig. 1 was found in at
least one species (Fig. 3). The most common trajector-
ies follow a continuous or sigmoid increase in crown
exposure with height, and often reflect the overall
community trajectory (Fig. 2d).
Most species (79%) have an intermediate crown
exposure at the juvenile stage (Table 1, Fig. 4) with the
remainder equally split between low and high relative
CE. At the adult stage most species have a relatively low
CE (57%). Only one species has a relatively low crown
exposure as both juvenile and adult (Gilbertiodendron
preussii, Fig. 3a), and this can be regarded as a whole-
life shade-tolerant species. One species has a relatively
high crown exposure as juvenile and adult (Aphanocalyx
microphyllus, Fig. 3i), and this can be regarded as a
whole-life shade-intolerant species. Many species (23%)
have intermediate crown exposure as juveniles and
adults (Fig. 3e), and simply follow the vertical light
gradient in the forest canopy. Nearly half of the species
(49%) start at intermediate levels and end up in rel-
atively low light conditions as adults (Fig. 3d). Three
species have relatively high CE as a juvenile, and a
Table 1 Crown exposure (CE) characteristics of 47 rain forest tree species. The species are ordered based on their maximal tree height. Adult stature (Hmax),
260
juvenile andet
L. Poorter adult
al. crown exposure, average changes in CE over the height range interval (CE change height-related) and over the monitoring period (CE
change temporal) and N are given. The Hmax is based on the 95th percentile of the population. Crown exposure is determined with a multinomial logistic
regression analysis for juvenile trees (5 m tall) and adult trees (average exposure at maximal tree height). For eight species (indicated with asterisks) the
smallest individuals were between 7.5 and 14 m tall. For these species the juvenile crown exposure has been extrapolated, based on the regression equations.
Letters following the CE indicate whether species have a relatively high (H), similar (Intermediate, I) or low (L) CE compared with other trees of the same
height class interval. The Nagelkerke r2 indicates how much of the variation is explained by the multinomial logistic regression model, and is given for the
significant regression lines. Note that insignificant regressions indicate that, on average, the CE is relatively constant over the height interval. Two species
are only known by their vernacular name. Nomenclature follows Jongkind (2004)

Species Family Hmax CE r2 CE change n

Height Temporal (y−1)

related
Juvenile Adult (m−1) Mean SE

Cola lateritia K. Schum. Sterculiaceae 13 1.48 I 1.88 I – 0.027 0.024 0.013 63

Craterispermum caudatum Hutchinson Rubiaceae 14 1.66 I 1.72 L – 0.003 − 0.007 0.003 411
Cola buntingii E.G. Baker f. Sterculiaceae 15 1.06 L 1.71 I 0.23 0.065 0.049 0.029 52
Baphia bancoensis Aubrév. Leguminosae-Pap. 17 1.36 I 2.10 I – 0.035 0.009 0.011 69
Diospyros sp. Ebenaceae 17 1.63 I 1.63 L – − 0.005 0.002 0.005 342
Syzygium gardneri Thw. Guttiferae 17 1.72 I 1.72 L 0.22 − 0.045 0.048 0.022 37
Garcinia afzelii Engl. Guttiferae 18 1.38 I 1.38 L – − 0.005 0.034 0.010 131
Rothmannia munsae (Schweinf. ex Hiern) Petit Rubiaceae 18 1.77 I 2.38 I – 0.040 0.006 0.004 40
Strephonema pseudocola A. Chev. Combretaceae 18 1.41 I 1.41 L – − 0.013 0.019 0.021 40
Drypetes sp. Euphorbiaceae 18 1.27 I 2.01 L – 0.025 0.008 0.006 56
Cola nitida (Vent.) Schott & Endl. Sterculiaceae 20 1.62 I 2.20 L – 0.023 − 0.011 0.008 91
Diospyros mannii Hiern. Ebenaceae 20 1.56 I 1.59 L – 0.001 0.028 0.006 404
Soyauxia sp. Medusandraceae 20 1.78 I 2.00 L 0.16 0.013 0.013 0.010 82
Memecylon lateriflorum (G. Don.) Bremek. Melastomataceae 21 1.53 I 1.81 L – 0.014 0.028 0.009 268
Mabagavi 22 1.50 I 2.56 L – 0.039 0.016 0.009 123
Placodiscus boya Aubrév. & Pellegr. Sapindaceae 22 1.81 I 1.81 L – − 0.019 0.027 0.013 53
Enantia polycarpa (DC) Engl. & Diels Annonaceae 22 1.43 I 2.21 I – 0.038 0.081 0.016 103
Chionanthus sp. Oleaceae 24 1.45 I 2.88 H 0.30 0.068 0.037 0.023 38
Diospyros gabonensis Gürke Ebenaceae 24 1.04 L 2.80 I 0.35 0.078 0.045 0.024 63
Carapa procera DC Meliaceae 24 1.04 L 3.00 H 0.71 0.094 0.025 0.019 39
Xylopia sp. Annonaceae 25 1.55 I 2.17 L 0.25 0.032 0.020 0.011 61
Calpocalyx brevibracteatus Harms Mimosaceae 25 1.53 I 2.57 L 0.09 0.028 0.017 0.009 100
Stachyothyrsus stapfiana (A. Chev.) J. Léonard & Leguminosae-Caes. 26 1.62 I 2.06 L – 0.018 0.122 0.019 92
Voorh.
Diospyros sanza-minika A. Cheval. Ebenaceae 27 1.41 I 2.52 L 0.07 0.038 0.042 0.004 934
Pausinystalia lane-poolei (Hutch.) ex Lane-Poole Rubiaceae 27 1.36 I 2.61 L 0.11 0.043 0.029 0.008 300
Trichoscypha aff.oba Aubrév. & Pellegr. Anacardiaceae 27 1.71 I 2.73 I – 0.041 0.023 0.014 69
Scytopetalum tieghemii (A. Cheval) Hutch. Scytopetalaceae 28 1.48 I 2.89 L 0.10 0.048 0.035 0.006 300
& Dalziel
Aphanocalyx microphyllus (Harms) Wieringa Leguminosae-Caes. 28 2.44* H 4.00 H 0.42 0.033 0.011 0.009 37
Mareya micranta (Benth.) Müll. Arg. Euphorbiaceae 29 2.16 H 2.77 L – 0.033 0.015 0.012 46
Panda oleosa Pierre Pandaceae 29 1.45 I 2.98 L – 0.049 0.038 0.011 54
Mae-I-Glu 30 1.84* I 3.19 I 0.59 0.066 0.051 0.020 36
Dacryodes klaineana (Pierre) H.J. Lam Burseraceae 31 1.80 I 2.60 L 0.06 0.027 0.055 0.013 147
Xylopia quintassii Engl. & Diels Annonaceae 32 1.34 I 3.56 H 0.39 0.092 0.068 0.017 71
Maranthes glabra (Oliver) Prance Chrysobalanaceae 32 1.75* I 3.11 I 0.29 0.052 0.030 0.012 47
Homalium smythei Hutch. & Dalziel Flacourtiaceae 33 1.29* I 2.81 L 0.31 0.054 0.044 0.021 49
Scottelia klaineana Pierre Flacourtiaceae 35 1.38 I 3.77 I 0.28 0.063 0.052 0.009 235
Maranthes aubrevillei (Pellegr.) Prance Chrysobalanaceae 35 1.31 I 3.29 L 0.39 0.065 0.028 0.009 77
Strombosia glaucescens J. Léonard Olacaceae 36 1.26 I 3.62 L 0.29 0.064 0.045 0.006 486
Calpocalyx aubrevillei Pellegr. Leguminosae-Mim. 38 1.25 L 4.14 I 0.48 0.078 0.027 0.005 187
Heritiera utilis (Sprague) Sprague Sterculiaceae 41 1.45 I 4.44 H 0.30 0.063 0.061 0.013 137
Dialium aubrevillei Pellegr. Leguminosae-Caes. 43 1.80* I 3.89 I 0.26 0.049 0.018 0.012 59
Gilbertiodendron preussii (Harms) J. Léonard Leguminosae-Caes. 46 1.23 L 3.58 L 0.55 0.058 0.023 0.006 213
Uapaca guineensis Müll.Arg. Euphorbiaceae 46 3.37* H 3.55 L 0.16 0.004 0.034 0.020 62
Anthonotha fragrans (Baker f.) Excell & Hillcoat Leguminosae-Caes. 48 1.28 I 4.35 I 0.46 0.076 0.018 0.009 64
Parkia bicolor (A. Cheval.) R. Capuron Leguminosae-Mim. 48 1.69* I 4.30 I 0.28 0.060 − 0.013 0.014 36
Nesogordonia papaverifera (A. Cheval.) Sterculiaceae 62 3.27 H 3.35 L – 0.002 0.052 0.014 43
Pterygota macrocarpa K. Schum. Sterculiaceae 63 2.25* H 4.15 I 0.45 0.048 0.028 0.014 48

© 2005 British
Ecological Society,
Journal of Ecology
93, 256–267
261 relatively low CE as an adult (Fig. 3g), although, in
Height–light absolute terms, this means that they remain in constant
trajectories of light levels. Only one species (Syzygium gardneri)
tropical tree species shows a significant absolute decrease in crown exposure
with an increase in height (Fig. 3d).
Fig. 2 Crown exposure index (CE) and average height–light
© 2005 British
trajectories for (a) all trees in the permanent sample plot
(n = 7460), (b) Gilbertiodendron preusii (n = 213), (c) Cola
Ecological Society,
buntingii (n = 52), and (d) all 47 species. The average height–
Journal of Ecology,
light trajectories and Nagelkerke r2 values are based on
93, 256–267 multinomial logistic regressions.
Species ranking in crown exposure of juveniles is to a
large extent maintained when trees increase in height.
Crown exposure is strongly correlated when 10 m and
20 m trees are compared (r = 0.68, P < 0.001, n = 43)
(Fig. 5a), with CE20 being consistently higher that CE 10
(delta CE = 0.42, paired t-test, t = 8.26, P < 0.001,
d.f. = 42). This correlation becomes weaker and
disappears when larger size classes are considered (e.g.
for 10 and 30 m tall trees r = 0.31, P = 0.09, n = 32; and
for 10 and 40 m tall trees r = − 0.16, P = 0.61, n = 12).
Between the juvenile stage and the adult stage species
crossovers in CE occurred in 564 out of 1081 possible
pairings (52%, Fig. 5b). The species with the most
extreme pairwise crossovers in CE are Anthonotha
fragrans and Calpocalyx aubrevillei, which both
have much greater CEs for adults than for juveniles
(Table 1).
-     
 
Species-specific slopes of CE against tree height varied
from −0.5 to 0.9 units CE per 10-m increase in height
(Table 1). Species also differed significantly (−0.01 to
0.12 units CE per year, Table 1) in temporal changes
over the monitoring period (, F46,6093 = 4.78,
P < 0.001, r2 = 0.36), with Stachyothyrsus stapfiana
showing an extremely large increase. As all its individ-
uals were found in a single 1-ha plot, and this plot
experienced significantly higher changes in CE
than the other 1-ha plots, we excluded this species from
further analysis with respect to temporal changes.
Although temporal and ontogenetic changes in CE
were not related to each other using a parametric cor-
relation (rp = 0.22, P = 0.14, n = 46), a non-parametric
correlation showed significance (rs = 0.30, P = 0.047,
n = 46).
Tree allometry appears to play a key role in getting
access to higher light levels. Species with slender
stems or narrow crowns realize larger changes in CE,
both height related (Fig. 6a,b) and temporally
(Fig. 6c,d).
 
Crown exposure is positively related to Hmax over a
large part of the vertical gradient (CE10 m height vs. Hmax,
r = 0.36, P < 0.05, n = 47; CE35 m height vs. Hmax, r = 0.68,
P < 0.001, n = 21, Fig. 7a). At larger heights the
relationship between crown exposure and Hmax disap-
pears, probably because of the small differences in
crown exposure, and small sample size (n ≤ 12). Adult
stature is closely linked to allometry, and height-
related changes and temporal changes in CE. Large-
stature species have more slender stems (r = 0.74,
P < 0.001, n = 42, Fig. 7b), a larger height-related
increase in CE (rs = 0.50, P < 0.001, n = 47) and a
larger temporal increase in CE (rs = 0.29, P < 0.05,
n = 46), as expected.
262
L. Poorter et al.

Fig. 3 Height–light trajectories of 47 rain forest tree species and the 95% confidence interval of the average vertical light profile
in the forest (dotted lines). The species are arranged in groups with similar changes in relative crown exposure between juvenile
(5 m tall) and adult stages (maximum height). Species are classified as occurring in relatively high light (above the 95% confidence
interval for the forest as a whole), intermediate light (within the confidence interval), or low light (below the confidence interval).
(a) Low-to-low light (n = 1), (b) low-to-intermediate light (n = 3), (c) low-to-high light (n = 1), (d) intermediate-to-low light
(n = 23), (e) intermediate-to-intermediate light (n = 11), (f) intermediate-to-high light (n = 3), (g) high-to-low light (n = 3), (h)
high-to-intermediate light (n = 1), (i) high-to-high light (n = 1). Different species are indicated by different symbols.

Discussion
We found evidence for all nine hypothesized light tra-
jectories, indicating that the pioneer–shade tolerant
dichotomy is far too simplistic, and that it is worthwhile
to look beyond the tolerances of seedlings and saplings.

    , 

  
At the stand level, the crown exposure increased in
a sigmoid fashion with the height of the trees. The
steepest changes in crown exposure occurred between
20 and 40 m, indicating the range where the light gain
may be large. Unfortunately however, little is known
about height growth dynamics of intermediate-sized
© 2005 British
Ecological Society, trees. Sterck & Bongers (1998) found that intermediate-
Journal of Ecology sized trees of 20 m height take large risks to get access to
93, 256–267 the canopy. They stretch out towards the canopy, and
Fig. 4 Relative commonness of different height–light trajec-
tories for 47 rain forest tree species. The relative light levels of
juveniles (below) and adults (above) are shown. Light levels
are classified as relatively low (CE lower than the average
forest level for that height), intermediate or relatively high.
The relative frequency of the light trajectory is indicated by
the thickness of the arrow, and the corresponding percentages.
263
Height–light
trajectories of
tropical tree species
Fig. 5 Changes in absolute crown exposure of species with
height. (a) Relationship between the crown exposure of species
at 10 m height (CE10) and 20 m height (CE20) (black circles),
and at 10 and 40 m height (CE40) (open circles). Pearson’s
correlation coefficient and significance levels are given. The
continuous line indicates the line y = x. (b) Average crown
exposures of juveniles and adults. Each line represents one
species. The relative position of several species changes with
size, indicating crossovers in light requirements among species.
© 2005 British
Ecological Society, Fig. 6 Relationship between height-related change in absolute CE and (a) stem slenderness, (b) crown diameter, and the
Journal of Ecology, relationship between temporal change in absolute CE and (c) stem slenderness and (d) crown diameter. Both the stem slenderness
93, 256–267 (i.e. height) and crown diameter of the species are averages for individuals with a d.b.h. of 15 cm.
have height-diameter ratios that are close to the ratios
under which they would buckle under their own mass.
Individual trees of a species occur under a wide
range of light environments (Fig. 2b, cf. Lieberman
et al. 1995; Bongers & Sterck 1998; Davies et al. 1998;
Poorter & Arets 2003). This has led, for example,
Lieberman et al. (1995) to conclude that ‘Instead of
high specificity and narrow tolerance we find broad
ecological amplitude; and rather than niche separa-
tion, we find a pattern of extensive overlap among the
great majority of the species in the assemblage’. The
question is whether high specificity and narrow tolerance
should be expected from long-lived, slow-responding
organisms such as trees growing in the dynamic envir-
onment as found in tropical rain forests. Mortality
rates among large trees are low, and the senescence
process is slow (Zagt & Werger 1998). Trees may take
several years before they show morphological responses
to the light environment in which they are growing
(Sterck et al. 1999), and light-demanding species may
therefore persist for several years in a low light environ-
ment before they die. At the same time canopy gaps are
formed at a regular rate. Gaps at the forest floor are
formed at a rate of 1% of the ground area per year (cf.
Jans et al. 1993), whereas gaps at 20 m height are formed
at a rate of 12% per year (Hubbell & Foster 1986). Over
time, individual seedlings and juveniles of shade-tolerant
species, especially long-lived ones, therefore have a
high chance of growing, at least for a while, in a canopy
gap. What counts from an evolutionary point of view,
though, is whether species differ in average population-
level light environments and we found that there were
264
L. Poorter et al.
Fig. 7 (a) Juvenile (filled circles) and adult (open circles)
crown exposure in relation to Hmax, and (b) stem slenderness in
relation to Hmax. The coefficients of determination and P levels
of each regression line are given.
indeed striking differences in height–light trajectories
(Fig. 3).
   
All nine of the light trajectories proposed for the
juvenile–adult transition (Fig. 1) were found in at least one
of our 47 species (Fig. 4), but only two strategies were
common: species that follow the forest light levels (and
thus the null model), and species that shift from follow-
ing the forest light levels when small to relatively low
levels when tall. The latter group contains typical sub-
canopy species that get stuck under other, taller, canopy
trees. Surprisingly few species complied with the classic
notion of whole-life shade tolerants (2% occurred con-
sistently in low light) and whole-life shade intolerants
(2% consistently in high light) (Figs 3 and 4). Extreme
switches are rare although three species started at rel-
atively high light levels and ended up at relatively low
ones, compared with the forest average (Fig. 5g). The
relative changes in Nesogordonia papaverifera and
Uapaca guineensis were particularly marked but, in
absolute terms, their crown exposure remained the
© 2005 British
same as height increased. Taller trees have a higher
Ecological Society, respiration load, and we might therefore expect that that
Journal of Ecology these species would approach their whole-plant light
93, 256–267 compensation point, and eventually die.
In absolute light terms only one species (Syzygium
gardneri) made a switch from high to low light (Fig. 3d).
Species that switch from high to low light between
the seedling and juvenile stage have been described
as ‘gamblers’ by Oldeman & van Dijk (1991), and as
‘cryptic pioneers’ by Hawthorne (1995). They may ger-
minate, establish and mature in the high light condi-
tions of gaps, before becoming overtopped by faster
growing pioneer species, but may then be able to persist
for a long time in the shade (Dalling et al. 2001). Ex-
amples of such species are Microdesmis puberula and
Myrianthus liberica in the Ivory Coast, Piper amalago in
Mexico (Oldeman & van Dijk 1991), and Alseis black-
iana in Panama (Dalling et al. 2001). Actually, many
more species may perceive a temporal decline in crown
exposure over time, either because they establish
in gaps and are overgrown in the building phase (Clark
& Clark 1992; Sterck et al. 1999), or because they
grow towards the canopy, get stuck in the shade of an
overtopping crown, and have to wait till this tree even-
tually dies off (Fig. 3g,i).
Species ranking in (absolute) crown exposure is to a
large extent maintained with height increase, probably
because most species follow the height-related null
model, especially over short height ranges (Fig. 3).
Over a larger height range, however, this consistency
in ranking gradually disappears. This indicates that
for some species the deviations from the null model
accumulate with size. When species approach the
canopy, individual light levels converge to similar
values. We conclude that, when species are compared
at two different height levels, rank reversals in irradi-
ance are not common but do occur, especially among
large-stature canopy species, but when species are
compared at two different life-history stages (juveniles
and adults) rank-reversals are commonplace (52% of
the cases).
-     
 
Twenty-four per cent of the individuals experienced a
change in absolute CE over the monitoring period. At
the population level, species show large temporal shifts
in average crown exposure. At first sight, the average
change in CE (−0.01 to 0.12 units CE year−1, Table 1)
might seem trivial, but if these rates are maintained
they may have important consequences for light parti-
tioning at reasonably short time-scales. If all species
start with a random distribution with respect to light,
and an average CE of 2, then the fastest declining spe-
cies will end up after 20 years with an average CE of
1.8, whereas the fastest increasing species will end up
with an average CE of 3.4. With species life spans that
exceed 100 years for most species such differences may
result in large crown exposure differences, especially
when growth rates are high. According to our expecta-
tion, temporal shifts in CE were positively correlated
with the height-dependent change in CE of the species.
265 Species that show a steep height-dependent increase in
Height–light CE also realize rapid temporal increases in CE. In con-
trajectories of trast to our expectations, however, temporal shifts in
tropical tree species CE were not related to the light requirements of the
species (Pearson’s r between temporal changes in CE
and CE5m = −0.04, P = 0.79, n = 47). High-light species
did not have a larger temporal shift in CE than low-light
species, probably because high-light species were already
concentrated in high-light environments by the juvenile
stage (Fig. 3g–i).
Species-specific differences in height–light trajector-
ies might be brought about by differences in low-light
mortality, height growth and allometry. Low-light
mortality has only a limited influence on the average
light trajectories of juvenile and adult trees (Poorter et al.,
unpublished data). This is in sharp contrast to the seed-
ling stage, where mortality rates provide an important
life-history filter, determining under what light condi-
tions species are found (cf. Li et al. 1996; Davies 2001;
Peña-Claros 2001; Montgomery & Chazdon 2002).
Unfortunately, we do not have data on height growth
rates of the trees. The importance of rapid vertical
expansion is underscored by the fact that species with
slender stems and narrow crowns realize both a faster
temporal and a faster height-related increase in crown
exposure (Fig. 6). By having little diameter growth and
limited lateral crown expansion, these species might be
able to invest more biomass in effective height exten-
sion, and take advantage of a steep vertical light gra-
dient. For these species the extension function of tree
architecture (an efficient height increase per unit bio-
mass invested) is more important than the light inter-
ception function (Poorter et al. 2003). They gamble on
reaping the future benefits (achieving a higher crown
exposure and attaining reproductive size), instead
of trying to enhance current light interception (King
1990; Kohyama & Hotta 1990; Sterck et al. 2001;
Kohyama et al. 2003).
Adult stature partly determines the light conditions
to which a species is eventually exposed. For instance,
tall species, by definition, grow to positions with high
light levels, in most cases having started from low light
when small. We expected therefore that the nine strat-
egies would differ in their adult stature and found that
that indeed was the case (, F8,38 = 4.2, P < 0.001).
We also found that large-stature species had higher
population-level CE (r = 0.93, P < 0.001, n = 47), and
that CE difference between juveniles and adults
increased with Hmax (Fig. 7a). Hmax is a simple indicator
of several population-level, allometric and biomechan-
ical characteristics and reflects differences in stem slender-
ness as well as in CE levels. This underscores the fact
that Hmax is an important life-history trait for which
tree species differentiate (Thomas 1996b; Thomas &
Bazzaz 1999; Turner 2001; Westoby et al. 2002). Adapting
stem allometry is an effective strategy for exploiting the
© 2005 British
Ecological Society, steep, vertical light gradient in the forest. Given this
Journal of Ecology, predictable and stable gradient, it is perhaps not sur-
93, 256–267 prising that evolutionary responses have led to a close
association between Hmax, light trajectories and allo-
metric traits.
We did not consider seed, seedling or sapling stages,
which are important phases in the life cycle of the tree,
where many changes occur. Many well-known com-
mercial species, such as the Entandrophragma and Khaya
species, may establish in the understorey and grow to
1 m height, but need a gap to recruit successfully to
larger size classes (Hawthorne 1995; Siepel et al. 2004).
We expect the inclusion of these stages to allow therefore
for the occurrence of more complex light trajectories,
and additional strategies (cf. Fig. 1). These diverse strat-
egies may help species coexistence in a tropical forest
environment where light availability is a critical factor
for survival and growth.
Acknowledgements
We are thankful to J. Poker and the GTZ for allowing
us to use the Grebo data collected by Hannsjörg Wöll
and Liberian colleagues, and to David Burslem, Peter
Grubb, Lawren Sack and an anonymous reviewer for
their enlightening comments. Douglas Sheil, Jerome
Chave, Agus Salim, Jerry Vanclay and William Haw-
thorne kindly allowed us to use their methodology
to analyse ontogenetic light trajectories. LP was sup-
ported by Veni grant 863.02.007 from the Netherlands
Organization of Scientific Research (NWO).
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Handling Editor: David Burslem

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Journal of Ecology,
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