POTENTIAL IMPACT OF CLIMATIC CHANGE ON TROPICAL RAIN

FOREST SEEDLINGS AND FOREST REGENERATION

T. C. WHITMORE
Geography Department, Cambridge University, Cambridge CB2 3EN, U.K.

Abstract. Tropical rain forests are dynamic and continually regenerating by growth of seedlings up
from the forest floor into canopy gaps that form on a cycle of usually a century of more in length.
Changes in seedling establishment, survival, and release in gaps could thus change canopy species
composition for a long time. Of likely climatic changes, evidence is presented that cyclone occurrence
and increased rainfall seasonality could have important effects on seedling ecology. These forests and
their species have lived through big Pleistocene and Holocene climatic changes, but today they are
fragmented by human impact and so have less resilience to future climatic change. Management to
accommodate climatic change should aim to reduce fragmentation and also canopy opening during
logging operations. These are the same practices as advocated for biodiversity conservation. Tropical
seasonal forests are also likely to be altered by expected climatic change, and also mainly at their
regeneration stage.

1. Introduction
Forests are dynamic. The canopy is in a continual state of flux, with gaps being
created by any one of many causes. Gaps range in size from a single tree crown to
many hectares, and soon become filled by regeneration. Tropical rain forests are
no exception to this general forest growth cycle. In them, gaps are usually filled by
seedlings that quickly grow up from a forest floor seedling bank to building-phase
then mature-phase canopy. Mature-phase canopy may persist, barring catastrophe,
for one to several centuries. What, then, is the interaction between forest regeneration, seedlings, and climate, and how might it change in a greenhouse world?
The germination and establishment of seedlings of many species in rain forests
is on the forest floor below the canopy. Their subsequent release occurs when
a canopy gap opens above them. All the evidence is that the main change at gap
creation is the 2040 fold increase in photosynthetically active radiation (PAR)
between closed forest (which receives c. 1 mol/m2 /day) and gap (a full clearing at
equatorial latitudes receives c. 35 mol/m2 /day). Some influences have been found of
water relations on seedling success, but so far little effect of plant mineral nutrients.
Canopy disturbance several centuries ago has been deduced in several different
rain forests from species composition and population structure today. In Central
America (Budowski, 1970) and Nigeria (Jones, 1955/1956), regrowth of forest
after past agriculture has been deduced, and in Sabah (Brown and Whitmore,
1992), after possibly past ENSO-related drought. On Kolombangara in the Solomon
Islands, either a cyclone or agriculture about a century ago is believed to have
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occurred (Whitmore, 1974). But eventually the signs of a particular disturbance

will disappear, because each tree generation reflects the circumstances that led to
its establishment. Thus the gap phase of the growth cycle is arguably the most
important because, once the gap has filled, species composition of that patch
remains the same for the rest of the cycle. The key question to address is how
climatic change might influence seedling success in gap-phase forest.
To discuss the potential impact of climatic change on seedling ecology is really
largely to map our ignorance. The sort of experimental work that would allow firm
predictions simply has not been undertaken. We can however make some guesses
as to what might happen, and in so doing indicate perhaps interesting topics for
future study.

2. Climatic Changes Likely to Influence Tropical Rain Forest Regeneration

Aspects of climatic change likely to be important to seedlings that emerge from
the analysis of Hulme and Viner (1998) are lower rainfall, greater seasonality, larger interannual differences, and lower soil moisture. These are all predicted to be
most marked for the two continental areas of rain forest, the Congo and Amazon
basins, and less marked for West Africa (where rain forest forms a narrow coastal
belt) and Southeast Asia (where it mainly occurs on islands). Only a rather small
increase in mean temperature is predicted. This is likely to be much less important for seedlings. Interestingly, ENSO events, which cause periodic drought and
have had considerable and well-documented effects in Indonesian and Malaysian
rain forests four times since 1980, seem either to show little change or to slightly
decrease. Cyclones, which are a major determinant of forest structure and composition (Whitmore, 1974), do not (according to Hulme and Viner, 1998) show major
changes, though Walsh and Pittock (1998) believes the cyclone belts might extend
polewards and that cyclone frequency could alter.
The changes Hulme and Viner predict develop progressively, and for a doubling
of atmospheric CO2 , the scenarios they map are reached some time between 2030
and the 22nd century. Thus, gradually, over a time scale of four decades at the
most rapid, various manifestations of greater periodic water shortage than today
develop.
We must remember that climates have fluctuated in the past (e.g., Zahn, 1994).
Tropical rain forests have expanded during the Holocene to attain their present
extent from a smaller area occupied at the last Glacial maximum (Flenley, 1998).
During the last Glacial to Interglacial transition period, tropical and subtropical
climatic fluctuations were rapid, with temperature sometimes dropping by 7 C
over a few decades, but they have been less rapid and severe since about 9000 BP
(Zahn, 1994).
Thus, many of the species that make up rain forests today have lived through
substantial climatic vicissitudes in the past.
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3. Rain Forest Seedling Sensitivity to Drought

3.1. OBSERVATIONS
There are two examples from Peninsular Malaysia of the importance of drought in
lowland evergreen dipterocarp rain forest. Shorea curtisii is a big emergent tree that
occurs gregariously on ridge crests of inland hills, especially on granite-derived
soils, and also on coastal hills. Burgess (1969) was able to show that these sites
have in common a propensity to drought, and develop very dry soil especially
in the early-year dry season. S. curtisii has seedlings that are better able than
those of other species to survive these conditions. Turner and Whitmore (1991),
working in Penang, north Malaya, discovered that the survival of seedlings of the
red meranti timber group of Shorea was lower in the annual dry spell than of
other dipterocarps. Northwards up the Malay Peninsula, the climate becomes more
seasonal, and early-year drought becomes more frequent and more marked. The
red meranti species group has its northern limit c. 100 km north of Penang. It seems
probable that inability of its seedlings to survive drought is the reason, especially
because the most northern location is a forest on deep, and therefore probably less
drought-prone, soils.
Another example comes from West Africa, where in Ghana there is a climatic gradient from perhumid climates in the southwest near Cote dIvoire, north
eastwards to more seasonal climates, and the forests change from evergreen to
semi-deciduous with a marked though not complete change in species composition. The high-rainfall soils in the southwest are more acid and have lower content
of plant mineral nutrients. In an experiment, nine species were grown together in
pots and subjected to a droughting regime equivalent to the main annual dry season
experienced by semi-deciduous forest. Ability to recover from drought mirrored
their natural occurrence along the climatic gradient: those from the more seasonal
places survived best, and the rain forest species did worst. Another experiment
showed little response to added mineral nutrients. It was concluded that water
availability in the dry season is more important than nutrient supply as a determinant of species occurrence (Veenendaal et al., 1994). Four separate experiments
on Malayan dipterocarp seedlings have failed to induce any response to added
nutrients (Turner et al., 1993; Burslem et al., 1995), perhaps because they were all
conducted at low light levels. The interaction of drought and nutrients on seedling
ecology has not been studied.
3.2. A CASE STUDY OF GAP PHASE REGENERATION
In the absence of any other direct evidence of the influence of water relations on
rain forest seedlings, let us look at a recent experiment on gap phase dynamics in a
dipterocarp evergreen rain forest and then consider how the response might change
in a drier climate.
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Figure 1. Differential survival of dipterocarp seedlings on the forest floor over 6 years, Danum,
Sabah. Seedlings of shade-tolerant species, in this case the two Hopea spp., characteristically survive
better in deep shade.

In this experiment (Brown and Whitmore, 1992; Whitmore and Brown, 1996),
artificial canopy gaps were made over natural mixed populations of seedlings of 11
different dipterocarp species in the Ulu Segama forest in Sabah, and response has
so far been monitored for 77 months. In the control, closed-forest plots amongst
two of the main species studied, survival is best of Hopea nervosa, a Medium
Hardwood species, and less in the Light Hardwood Shorea johorensis (Figure 1).
Because of this property, which is well known in dipterocarps, H. nervosa is
described as a shade-bearer and S. johorensis as a light-demander. Other Light
Hardwood species showed similar survival to S. johorensis and H. beccariana,
another Medium hardwood species, is similar to H. nervosa (Figure 1).
The nine different gaps that were created were grouped into size classes by
the mean daily amount of PAR received at their centers tiny (c. 4 mol/m 2 /day),
small (c. 10), medium (c. 14), and large (c. 19). By 53 months (Figure 2a), there
had been little growth in the control plots (c. 1 mol/m2 /day), H. nervosa had
grown by about the same amount in all gaps but in small gaps (10 mol/m2 /day),
and larger S. johorensis had grown progressively taller with greater gap size. By
77 months (Figure 2b), H. nervosa still showed little growth and S. johorensis
showed an even stronger difference. This experiment shows that S. johorensis is
able to utilise extra PAR, received either with longer time or in a larger gap, for
height growth, but H. nervosa is not. S. johorensis can be described as a lightdemander and H. nervosa as light-indifferent. The other species monitored were
individually less common. Collectively, the Light Hardwood species behaved like
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S. johorensis and the Medium Hardwoods like H. nervosa. The exception was
Parashorea malaanonan, also shown on Figure 2. This is a Light Hardwood that
puts on little height because of very severe apical damage by herbivory.
We believe this experiment shows the means whereby light-demanding Light
Hardwood rain forest species are favored in silviculture by big canopy gaps whilst,
if a forester wants to favor so-called shade-tolerant Medium Hardwoods, he makes
only tiny gaps. The experiment also shows the very important role that can be
played by species-specific herbivory (Coley, 1998).
The question of interest here is how the dynamics might be affected by the
greater seasonality and drought arising from climatic change.
The seedling bank below closed canopy is replenished with fresh fruits. Dipterocarps are triggered to flower by a stronger than usual dry season and do so heavily
once every 47 years. Greater seasonality would perhaps trigger more frequent
mass fruiting (Borchert, 1998), hence more frequent seedling bank augmentation.
This could lead to the long persistent, shade-tolerant Medium and Heavy Hardwood
species losing their height advantage so that even in small gaps Light Hardwoods
become their equal. But dipterocarps do not fruit heavily if two dry years occur in
succession. Their phenological pattern would probably become totally disrupted
were strong dry seasons to become annual.
Drier climate and drier soil could differentially affect dipterocarp seedling establishment and survival. Indeed, the evidence of Shorea curtisii and other red merantis
reviewed above suggests it would. These are Light Hardwoods and behave in gaps
like S. johorensis in the experiment. Herbivory patterns might change. Overall,
the balance of species in the seedling bank might therefore alter, perhaps to the
disadvantage of red meranti Shorea, which are commercially the most important
group.
Thus, in general terms, the differential seedling release in gaps we have revealed
in the Sabah experiment would probably be altered if the climate became drier and
more seasonal. We have no means without an elaborate droughting experiment
conducted over several years within the forest to predict which species or species
groups might be favored. Long-term monitoring of seedling demography along the
climatic gradient northwards through Peninsular Malaysia into southern Thailand
could also be revealing.
3.3. CONCLUSIONS
These observations and the experiment provide straws in the wind which indicate
that regeneration would probably change were the rain forest climate to become
more seasonal. The change would be slow because gap-phase regeneration in
primary forest commonly takes place once a century or less.
It must also be borne in mind that the postulated rate of change is similar to
the various fluctuations that have occurred during the last Glacial to Interglacial
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Figure 2. Dipterocarp seedling growth in canopy gaps of different size (a) after 53 months, (b) after
77 months (Whitmore and Brown, 1996, Figure 2).

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transition period, and less severe than the big change that has occurred in much of
the range of tropical rain forests since the last Glacial maximum (and earlier ones).
Before human intervention, species are likely to have changed their ranges as
the places where their seedlings could be successful slowly moved in concert with
climatic fluctuation. The major difference today is that human-induced climatic
change affects forests that are themselves altered by humans.

4. Impact of Climatic Change on Forests under Anthropogenic Stress

In much of the tropics, rain forests are being cleared and fragmented to make
way for agriculture. They increasingly persist as islands set in a sea of plantations
or farms. Many of the big blocks are suffering attrition at their edges. This was
quantified by FAO in its 1990 assessment of the state of the worlds forests by
computing the ratio of perimeter to area, and the amount of edge (defined as a
10 km broad band) to core (FAO, 1993).
It is likely that near the edge, forest interior microclimate will be altered (Malcolm, 1998), though we so far have data only from one location in the Amazon
(Kapos, 1989). This could alter gap-phase regeneration. At the Amazon site, small
fragments and edges have suffered greater windthrow and this too may influence
regeneration.
The additional interaction between loss plus fragmentation and climatic change
is that the possibility of the slow movement of species across a landscape to track
the changing position of suitable regeneration niches is blocked.
A second major contemporary human impact on tropical rain forests is mechanized timber removal. This is particularly extensive in Asia, which supplies most of
the tropical hardwoods entering international trade. By early next century there will
be virtually no unlogged Asian rain forest outside conservation areas. Logging does
not destroy forests, but during logging the canopy is opened to a greater extent than
occurs naturally and as a consequence the species composition of both plants and
animals in the regrowth is more biased to light-demanding, disturbance-favoring
species. Desiccation could become of biological importance if strong dry seasons
occurred more frequently during or soon after a logging operation. After logging
there is much dead wood and litter and the forest may catch fire in dry weather
(Goldammer and Price, 1998). This has recently occurred twice in the Malesian
dipterocarp forests. In 1983 4.5 million ha were burned or damaged by drought in
Borneo (Lennertz and Panzer, 1984).
Thus, both forest fragmentation and logging exacerbate the effects of climatic
change by allowing the forest floor to desiccate more strongly and this may have an
effect on species composition of the regenerating forest. Secondly, fragmentation
itself may inhibit or prevent the movement of species to suitable regeneration niches, thus closing the means whereby species have adapted to previous fluctuations
in climate.
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5. Implications for Management

Human destruction and disturbance of tropical rain forests is more rapid and
dramatic than that likely to be caused by the predicted changes to equatorial
climates over the next century or so. It follows that the main focus for forest
and biodiversity conservation has to be on these direct impacts. A start has been
made with the imposition by ITTO (International Tropical Timber Organisation)
of the year 2000 as the target date by which all forests will be managed for timber
production in a sustainable manner, by following published guidelines (ITTO,
1990). The same practices which minimize logging damage will also make the
forest less susceptible to alteration as the climate becomes slightly warmer and more
seasonal. These practices are well known. Overall they aim to minimize canopy
opening. This means logging should be on polycyclic systems, extracting just a
few large mature trees per hectare, and causing minimum damage by following
well-known rules (marking of trees for harvest, directional felling, preplanned
well-routed extraction tracks, using winches and cables not tractors to pull logs to
the tracks). The Amazon and Congo basins are the regions whose climate is likely
to change most, and are therefore in need of greatest care.
In addition, good land use planning is needed to ensure that forest remains in
large blocks in which production forest lies contiguous with protection forest. Small
fragments should be linked to each other and to large blocks in order to maintain
large populations of individuals in the short term, and to allow the possibility of
migration in the longer term.
All these measures can be summarized as building for resilience. It can be
seen that the practices which will mitigate effects of climatic change are also
those recommended for forest maintenance and biodiversity conservation in an
increasingly human-dominated landscape.

6. Tropical Seasonal Forests

Less is known about seedling ecology in tropical seasonal forests. Fire is a major
factor. Much regeneration is by coppice. Human impact has been more extensive
and severe than on rain forests, and tropical forests are already much more reduced
and fragmented than rain forests. Hulme and Viner (1998) predict similar warmer
and more seasonal trends for the seasonal tropics except for a swathe from eastern
and northen Africa across to India, where rainfall could be as much as 50 percent
higher and seasonality reduced. It can be predicted that in seasonal forests, as in
rain forests, species composition will slowly be transformed because of altered
conditions for seedling establishment due to climatic change, either directly or
via altered fire regimes. However, vegetative regrowth from stumps will have a
powerful buffering effect. The severe fragmentation of natural and semi-natural
seasonal forests makes species movement difficult and perhaps species extinction
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from this cause will be greater than in rain forests. As in rain forests, species loss
due to climatic change will, I predict, be trivial compared with that caused by more
direct human action.
Acknowledgements
Thanks are due to WWF for enabling participation in this workshop and to N. D.
Brown for the data and its analysis in Figure 1.
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