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RT Corlett, Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Menglun, Yunnan, China
© 2018 Elsevier Inc. All rights reserved.
Tropical rainforests are the tall, dense, multilayered, broad-leaved evergreen forests that form the natural vegetation in those areas of
the tropics where the climate is always hot and the dry season is brief or nonexistent (Corlett and Primack, 2011). Forests that
receive <1800–2000 mm of rainfall annually or experience a dry season longer than 2–3 months usually have an increasing
proportion of deciduous trees. Evergreen rainforests may also extend into areas with lower rainfall where groundwater is available,
such as along rivers, or where dry season water stress is reduced by fog or overcast skies or, near the margins of the tropics, by cool
winters. On tropical mountains, above around 1000–1200 m elevation, and north and south of the tropics, lower temperatures
result in forests that are evergreen, but have a lower canopy height and less complex structure. These forests are usually referred to as
“montane” and “subtropical.” Note that local usage of the term “rainforest” is not standardized across the tropics, so care needs to
be taken when comparing forests in different regions.
Until recently, tropical rainforests formed a broad belt around the equator, interrupted only by the oceans, by dry climates in eastern
Africa and northeastern Brazil, and by the Andes Mountains in South America. There were also extensions north and south of this
belt to the margins of the tropics wherever rainfall was sufficient, in Central America, along the southeast coast of Brazil and the east
coasts of Madagascar and Australia, and in Southeast Asia, northeast India, and along the crest of the Western Ghats in western
India. Smaller areas of tropical rainforest occur on some tropical islands, usually where additional rainfall is produced by the uplift
of the prevailing winds as they blow over mountains.
The oceans are a highly effective barrier to most forest organisms so the floras and faunas of the different rainforest regions have
evolved independently for tens of millions of years. Mountains and dry areas can also isolate rainforests, but these barriers are
generally less permanent than ocean basins. The shallower seas that separate the islands of Southeast Asia and separate New Guinea
from Australia were only intermittent barriers, since the low sea levels during the glacial periods created dryland connections many
times in the last 2 million years. It is therefore useful to divide the world’s tropical rainforests into five major regions, each with its
own distinctive fauna and, to a somewhat lesser extent, flora (Corlett and Primack, 2011):
1. Neotropics (South and Central America, with small areas on some Caribbean islands)
2. Africa (Central and West Africa, with outlying islands of rainforest in East Africa)
3. Madagascar (along the east coast)
4. Asia (much of Southeast Asia, extending into northeast India, with outlying areas in the Western Ghats of India and southwest
Sri Lanka)
5. New Guinea and Australia (most of lowland New Guinea and small areas in northeastern Australia)
Tropical rainforests are of vital global importance for two major reasons: their exceptional biodiversity and their key role in global
carbon cycles. An estimated two-thirds of flowering plant species grow in the tropics (Pimm and Joppa, 2015), with the highest
diversities in rainforests. Similar patterns are shown by birds, mammals, and amphibians, as well as most invertebrate groups that
have been assessed (Corlett, 2014a). Tropical forests – mostly rainforests – store around a quarter of the global terrestrial carbon
pool and they account for a third of net primary production. Thus, they have a major influence on the exchange of carbon between
the land and the atmosphere. Moreover, these forests are currently a large sink for the carbon dioxide produced by human activities.
Note, however, that these two key global roles of tropical rainforests are determined by different components of the total biota: their
biodiversity reflects the vast numbers of rare plant and animal species in these forests, while most of the carbon passes through a few
common species of giant trees (Fauset et al., 2015).
Tropical rainforests are also important for local people and make major contributions to the economies of the countries in which
they occur. They are a potentially sustainable source for a wide range of products, including timber, firewood, food, medicines, and
other plant products, as well as services, including local climate amelioration, slope protection, and the provision of clean water.
Timber production is a major source of income and employment in tropical rainforest countries and many tropical cities depend on
forested catchment areas for their water supply. Tropical rainforests have also served as a “land bank” for the development of
commercial agriculture in many countries. This is obviously not a sustainable use, but it has contributed to the spectacular
economic development in some of these countries over the last few decades.
Tropical rainforests similar in appearance to those that are found today have existed for 60–70 million years, although their floras
and faunas have changed over time. The extent of these forests has expanded and contracted, with major extensions toward high
latitudes during the warmest, wettest periods and major retractions to equatorial regions during cooler, drier ones. The changes in
rainforest area and distribution were particularly dramatic during the ice ages (glacial periods) of the last 2.5 million years, with
lower temperatures and rainfall excluding tropical rainforests from many areas that they occupy today, while lower sea levels
permitted the expansion of rainforests elsewhere, particularly in Southeast Asia (Corlett, 2014a). By around 9000 years ago, climatic
conditions in the tropics were broadly similar to the present day, although fluctuations continued and even the wettest areas seem to
have experienced occasional severe droughts.
Intact tropical rainforests appear to be resilient to natural climatic variation, but accelerating development globally and across
the tropics in the last 50–60 years has put increasing strains on this capacity. Global climate change, resulting largely from
anthropogenic greenhouse gas emissions, has raised temperatures across the tropics by around 0.7–0.8 C since 1950 (IPCC,
2013). It has probably also changed rainfall patterns, but the large variation between years in the tropics makes this difficult to
detect with any certainty. The impacts of these changes have so far been unclear, except on tropical mountains, where plants, birds,
and insects have moved upslope in response to rising temperatures (Freeman and Freeman, 2014).
Human activities have also changed climates on a local and regional scale in the tropics. When large, continuous, tracts of
rainforest are fragmented by crops or pasture, the remaining forest patches tend to dry out and become susceptible to fires that are
started in the open areas. Even if the forest is not cleared, logging – the harvesting of trees for timber – opens up the canopy and
increases the risk of fire. Moreover, rainforest trees, with their larger leaf area and deeper roots, transpire more water than crops,
pasture, or bare soil, cooling and humidifying the atmosphere, so when forest is cleared, the regional climate becomes warmer and
drier (Devaraju et al., 2015). The combined result of these impacts, coupled with the expanded use of fire as a tool for forest
clearance and land management, has been a huge increase in the frequency and extent of fires in rainforest regions, particularly in
the dry years that occur at irregular intervals. These fires in turn lead to regional haze episodes that are hazardous to human health
and aviation and may further suppress rainfall (Marlier et al., 2014).
Predictions for future climate change depend on complex global models, driven by assumptions about how greenhouse gas
emissions and other climate drivers will change in the future. These models predict an additional 1–2 C warming in the tropics
by 2050 and 2–5 C by 2100, depending on the assumptions (IPCC, 2013). The biggest uncertainties concern the future trajectory of
greenhouse gas emissions, since this will be influenced by socioeconomic factors, technological change, and political will. Ongoing
global negotiations on reducing emissions aim to keep warming below 2 C, but achieving this will require larger reductions than
currently seem likely. Even with the most optimistic assumptions, tropical temperatures by 2100 are predicted to be the warmest
that they have been for several million years. Predictions for tropical rainfall vary considerably among models, but the general
pattern is a moderate increase in rainfall in most, but not all, areas. Predictions for climatic extremes vary even more, but an
increased frequency of severe droughts and an intensification of cyclones are both supported by some models. Two major droughts
occurred in the last decade in Amazonia. Even if the frequency and duration of droughts does not change in terms of rainfall, higher
temperatures will increase the drought stress experienced by plants.
We know that tropical rainforests have survived past changes in climate, including the 4–5 C temperature swings over the last
2.5 million years. However, these past changes were considerably slower than those expected over the next few decades, and
temperatures were usually cooler than today. Moreover, in contrast to the past, today’s rainforests are fragmented and subject
to multiple additional stresses, including logging, hunting, and fires. Predicting the impacts of future climate change on
tropical rainforests is difficult because there is no analogue for the warmer climates expected for the second half of the century,
either today or in the last million years or so. These “no-analogue futures” mean that predictions must be based on models
and experiments. The models cannot be tested by comparisons with observations, so the accuracy of their predictions depends
on how well they incorporate the key processes involved. Experiments are preferable since all relevant variables can be
controlled, but in tropical rainforests, these have been limited by the sheer size of rainforest trees, which makes any form
of enclosure impractical.
Temperature
Although predictions of rising temperatures throughout the tropics are scientifically robust, with the only disagreements being
about the rate of warming, the potential impacts of this warming on tropical rainforests are still poorly understood (Corlett, 2011).
Tropical rainforests are currently excluded from many land areas by drought or cold, but they are not excluded from any area by high
Tropical Rainforests and Climate Change 27
temperatures, so the upper limit of their thermal tolerance is unknown. The predicted 2–5 C temperature increase may not sound
much, but it is large relative to present variability. A temperate-zone tree may have to survive a temperature range of 50 C or more
during its lifetime, while a tree in equatorial rainforest may spend its entire life in a 22–34 C range.
We know an increasing amount about how rainforest insects, frogs, and tree seedlings respond to warming under
controlled conditions, but nobody has yet warmed up an adult rainforest tree or, indeed, an entire rainforest. At one extreme
are suggestions that rising temperatures will lead to a drastic “lowland biotic attrition,” with species dying or retreating
upslope as their temperature tolerances are exceeded, while others suggest that most species will be able to adjust to a
moderate rise in temperature so there will be little change overall (Corlett, 2011). This adjustment could occur by acclimation
(physiological or behavioral adjustment during the lifetime of an individual organism) or adaptation (genetic change as a
result of natural selection over multiple generations). Given the projected rates of warming over the remainder of this century,
significant genetic adaptation will only be possible for short-lived species that will go through many generations in this time
and then only if there is heritable variation in thermal tolerance. Acclimation is the only option for long-lived organisms, like
elephants and large trees. Unfortunately, neither of these processes is well understood at present for tropical rainforest
organisms, so it is not yet possible to narrow down the range of possible futures. Moreover, increasing temperatures will be
accompanied by an increased frequency and intensity of extreme heat events, and these may have a very different impact from
increases in mean temperature.
Rainfall
Although the climate models differ greatly in their predictions for future rainfall in the tropics, the potential impacts of any changes
in rainfall on tropical rainforests are better understood than those of temperature. This is partly because rainfall varies a lot more
among years than temperature, so it is possible to use dry (or wet) years as a model for a drier (or wetter) future, but also because
rainfall exclusion experiments – where part of the rainfall is diverted before it reaches the ground by plastic sheets and gutters – are
much easier to do in intact rainforests than experimental warming would be. Both observations and experiments show that
exceptional droughts kill trees, particularly big ones, but it is still not clear how they do this. There is evidence for both hydraulic
failure – a breakdown of water transport in the xylem due to the formation of air bubbles – and carbon starvation as a result of
closure of the stomata. Real droughts – but not rainfall exclusion experiments – are often accompanied by clear skies, higher
temperatures, and lower humidities, which can all exacerbate the impact of drought alone.
Short-term droughts kill individual trees, but the forest can recover in years of normal rainfall. Multiple droughts and long-term
drying trends, in contrast, are expected to lead to changes in species composition, with the more drought-tolerant species being
favored. This will make the forest more resilient to the impacts of future droughts, but there is also a risk that an increase in fire
frequency resulting from other human impacts will interact with the drying climate to produce an abrupt switch from rainforest to
grassland or savanna.
Carbon Dioxide
Rising carbon dioxide levels not only are the biggest single driver of climate change but also have a direct impact on plant
physiology and growth. In particular, higher CO2 levels can enable plants to fix more carbon for the same amount of water loss
through their stomata. In both models and experiments, increases in carbon dioxide can increase photosynthesis and water-use
efficiency: the so-called CO2 fertilization effect (Schimel et al., 2015). Modeled futures for tropical rainforests look a great deal more
optimistic when this effect is included. In particular, droughts have far less impact on rainforest productivity and long-term
persistence than they do if the impact of climate change alone is modeled. There is some evidence that CO2 fertilization is already
having an impact on tropical rainforests, but there is currently no agreement on the potential longer-term influence.
If photosynthesis is limited by nutrient shortages, as is likely on typically nutrient-poor rainforest soils, or by rising temperature,
then plants will not be able to make use of extra CO2.
fuel consumption. Predicting human behavior is inherently difficult, because of the massive adaptation potential of individuals and
societies, but these indirect impacts could potentially be more significant than the direct ones.
Adaptation
At least 2 C of warming in the tropics now seems inevitable and more than this is likely. This means that, in addition to mitigation,
we must also be concerned with climate change adaptation, that is, reducing the vulnerability of tropical rainforests to climate
change. Adaptation is not an alternative to mitigation, since business-as-usual climate change will eventually exceed any plausible
capacity to adapt, but it is a necessary precaution. Moreover, reducing the vulnerability of rainforests to climate change will also
benefit the mitigation efforts described in the previous section.
Two general mechanisms have been suggested for vulnerability reduction: reducing nonclimate stresses and increasing connec-
tivity across climate gradients. The most important nonclimate stresses on tropical rainforests are forest clearance, fragmentation,
and logging, which tend to dry out the remaining forest and greatly increase the risk of fires. These activities also bring economic and
livelihood benefits, so banning them may be impractical, but it may be possible to manage clearance in a way that leaves large
continuous blocks, rather than isolated patches, and to reduce logging damage by the adoption of reduced-impact logging
guidelines. Connectivity across climatic gradients, such as those associated with altitude, is important because it allows rainforest
species to move in response to climate change, for example, upslope in response to warming (Corlett, 2015). Although rainforest
trees can move only during the seed dispersal phase, once a generation, the steep temperature gradients in rugged topography –
roughly 0.6 C for every 100 m increase in elevation – mean that even a moderate dispersal capacity can be useful. However, the land
area available usually decreases with elevation and mountain summits set an upper limit on movement, so the potential for
adaptation by movement is limited.
The capacity for climate change mitigation in tropical rainforest regions is significant, but relatively small in comparison with total
global greenhouse gas emissions. These are dominated by emissions from fossil fuel combustion, particularly in China, the United
States, and Europe. If the protection of tropical rainforests is a global priority, as their role in both carbon sequestration and
biodiversity conservation suggests it should be, then this protection must include a global agreement to reduce fossil fuel emissions.
While nobody expects the fate of tropical rainforests to overshadow the numerous other factors that influence climate negotiations,
this fate is both important in itself and symptomatic of broader problems in the relationship between humans and the
natural world.
References
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Corlett RT (2015) Plant movements in response to rapid climate change. In: Peh KS-H, Corlett RT, and Bergeron Y (eds.) The Routledge handbook of forest ecology, pp. 517–526.
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Tropical Rainforests and Climate Change 29
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