Restoration of Degraded Tropical Forest Landscapes

David Lamb et al.

Science 310, 1628 (2005);
DOI: 10.1126/science.1111773

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 REVIEW
Restoration of Degraded Tropical
Forest Landscapes
David Lamb,1* Peter D. Erskine,1 John A. Parrotta2

tive forests are still present in the landscape.

The current scale of deforestation in tropical regions and the large areas of degraded Well-documented examples where natural re-
lands now present underscore the urgent need for interventions to restore biodiversity, generation has occurred over very large areas
ecological functioning, and the supply of goods and ecological services previously used are Puerto Rico (3), Tanzania (4), Costa Rica
by poor rural communities. Traditional timber plantations have supplied some goods (5), and Brazil (6). Species-rich forests can
but have made only minor contributions to fulfilling most of these other objectives. develop in this way, but such forests often
New approaches to reforestation are now emerging, with potential for both overcoming contain only a subset of the original plant

Downloaded from www.sciencemag.org on August 30, 2013

forest degradation and addressing rural poverty. or wildlife species (3, 7). Although it is rarely
possible to determine the proportions or iden-

O
ne of the defining events of the past goods (e.g., timbers, medicines, and foods) tities of missing plant species, the most com-
century was the astonishingly rapid once provided by the original forests to the mon absentees are the large-fruited plant
decline in the extent of tropical people living in these areas. species because of the absence of appropriate
forests. An estimated 350 million hectares Neither agricultural development nor past dispersal agents. Foresters have sought to
have been deforested, and another 500 million forms of reforestation have been sufficient increase the populations of commercially im-
hectares of secondary and primary tropical to provide sustainable livelihoods and envi- portant timber species in such secondary forest
forests have been degraded (1). The damaging ronmental services over the large areas of by enrichment planting (planting target species
consequences of this include the loss of degraded land that have developed. Despite under canopy gaps or along cleared strips) (Fig.
ecological services (such as biodiversity and the expansion of protected area networks, 2). The same technique might be used to
watershed protection), the loss of many goods there has been an overall gradual simplifica- improve biodiversity by adding species that
(such as timber and nontimber forest products), tion and homogenization of some of the are otherwise unable to colonize and regenerate
and the loss of means of existence for forest- world_s most biologically diverse landscapes. or are ecologically threatened or vulnerable.
dwelling people. These losses have fallen par- It is unclear just what the long-term conse- Natural recovery of degraded forest areas
ticularly heavily on the rural poor in tropical quences of this might be. is not inevitable, and recovery is difficult
countries, where the livelihoods of at least 300 In recent years, new forms of reforestation where the system has crossed an ecological
million people now depend upon these degraded have been tested that may offer additional threshold and reached a new steady state con-
or secondary forests (1). ways of dealing with degraded tropical forest
Until recently there were three major re- landscapes. These include improvements in the
sponses to this process of forest degradation. management of secondary or regrowth forests
One was to expand networks of protected areas as well as more complex forms of reforestation
to help protect the remaining biodiversity. In where forest cover has been entirely lost
this response, the focus has largely been on ESupporting Online Material (SOM) Text^.
making the selection of candidate sites as There is clear evidence that biodiversity
representative and comprehensive as possible conservation can be enhanced by the careful
(2). A second was to improve agricultural pro- location of protected areas (2). Likewise, im-
ductivity on abandoned lands in order to proved methods of regrowth management and
improve the livelihoods of communities living reforestation should also help restore bio-
in these areas. The third approach has been to diversity to degraded landscapes (Fig. 1).
undertake some form of reforestation. Much of
Accelerating Natural Recovery Fig. 1. A conceptual diagram of changes in forest
this has been done with the use of industrial cover in a landscape over time as a consequence
monocultures involving a limited number of One way of increasing forest cover is to protect of agricultural intensification. Forest cover is
species from a remarkably small number of and manage the large areas of secondary or shown as a solid line, and the corresponding
genera (particularly Pinus, Eucalyptus, and regrowth forests now present. Not all degraded change in biodiversity is shown as a dotted line.
Acacia). Although many of these plantations lands are completely deforested, and they vary Biodiversity loss occurs as forest cover declines,
although the magnitude of this loss depends on
have been productive and generated goods in forest cover, degree of fragmentation, and
the extent and location of the protected area
such as pulpwood, few provide the variety of extent to which biodiversity has been lost. They network. When reforestation begins to occur
also vary in their capacity to recover unaided if (shaded area), it will increase forest cover, but
1
Rainforest Cooperative Research Center and School further disturbances can be prevented. Succes- any corresponding improvement in biodiversity
of Integrative Biology, University of Queensland, sional development (or self-repair) can be rapid depends on the types of reforestation carried out.
Brisbane 4072, Australia. 2Research and Develop- Trend A depicts a scenario where secondary forest
ment, Forest Service, U.S. Department of Agriculture,
at sites where forest clearance has occurred
is protected and where connectivity is enhanced
4th floor, RP-C, 1601 North Kent Street, Arlington, relatively recently (years versus decades); where
by reforestation using a diverse range of native
VA 22209, USA. some residual trees, seedling banks, and soil species; trend B, where reforestation relies solely
*To whom correspondence should be addressed. seed stores composed of native species re- on extensive monoculture plantations of fast-
E-mail: [email protected] main; and where intact, biodiversity-rich na- growing exotic species.

1628 9 DECEMBER 2005 VOL 310 SCIENCE www.sciencemag.org

 REVIEW
dition (8, 9). A common example is when deg- ability of additional native species to reach the the undisturbed forest in its tree species com-
radation leads to topsoil loss and a reduction site from nearby intact forest, principally position, although structural recovery is still
in soil fertility, complicating recolonization of through seed dispersal by frugivorous birds incomplete (18). The approach allows key
these sites for many of the original species. and mammals. In such cases, small-fruited species to be targeted (e.g., large-fruited spe-
Another threshold is commonly crossed when species are generally more likely to colonize cies) but requires sufficient ecological knowl-
sites become occupied by grasses. This than large-fruited species. The approach also edge to be able to collect seeds and germinate
increases the risk of wildfires, particularly in runs the risk of facilitating colonization by large numbers of seedlings from a wide variety
the seasonal tropics, which then reduces woody undesired weed species. of species.
plant recruitment and favors the further spread The other approach uses a much greater The key limitation to the use of such res-
of grasslands. There are many examples number of species representative of more toration plantings is their high cost (20). They
throughout the tropics of extensive grasslands mature successional stages and bypasses the do not supply significant volumes of com-
that persist over time despite being entirely natural successional sequence. Plantings are mercially useful goods such as timber, and
surrounded by forests (10). Therefore, usually there are only limited markets for
even though natural regeneration is po- the ecological services they provide.
tentially the cheapest way of fostering re- Hence, restoration plantings are probably
forestation over large areas, it is also the an option that can only be used in a
riskiest option because thresholds may relatively small number of situations and
have been crossed or because excluding rarely in the most severely degraded
further disturbances is difficult (9). tropical landscapes, except where the

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potential environmental benefits or costs
Plantings and Plantations of inaction (as in mined land or man-
Most deliberate efforts to overcome grove restoration) may justify the
degradation involve tree planting. How- required investment (SOM Text).
ever, even traditional forms of timber Plantation establishment to provide
plantation can be risky operations, and, goods and ecological services. An alter-
where species selection or early stand native is to establish plantations that
management are inappropriate, plan- provide goods for which there is a market,
tations can fail (11, 12). Planting to gen- such as timbers, but also generate a larger
erate ecological services as well as goods range of ecological services than the more
is even more difficult, because trade-offs Fig. 2. It is difficult to develop restoration methods at a traditional industrial timber plantations.
must be made between the productivity particular site that optimize financial and livelihood This approach seeks a balance between
benefits as well as generate improvements in biodiversity
of desired goods (i.e., timber) and provi- (top right corner). Traditional monoculture plantations of the financial benefits of industrial timber
sion of ecological services (i.e., bio- exotic species (arrow 1) mostly generate just financial plantations (which enable large areas to
diversity) and the techniques to achieve benefits, whereas restoration using methods that maxi- be reforested) and the biodiversity gains
these simultaneous goals are still being mize diversity and (arrow 2) enhance biodiversity yields possible from carrying out a more com-
developed. Some of the approaches are few direct financial benefits to landowners, at least in the plete ecological restoration.
summarized in Table 1, which also shows short term. Protecting forest regrowth (arrow 3) generates The simplest way this might be done is
improvements in both biodiversity and livelihoods,
their capacity to supply services as well although the magnitude of the benefits depends on the to use monocultures of native species
as goods. The choice of approach will population density of commercially or socially important (particularly those that are fleshy-fruited
depend on the socioeconomic circum- species; these can be increased by enrichment of or have propagules dispersed by forest
stances of the land owner as well as on secondary forest with commercially attractive species wildlife) rather than exotics. Although the
the ecological situation (soil fertility, (arrow 4). Restoration in landscapes where poverty is biodiversity gains are modest, they are
extent to which natural forest remnants common necessitates attempting both objectives simul- more likely to create environmental
taneously. But, in many situations, it may be necessary to
remain in the landscape, etc.) (13). give initial priority to forms of reforestation that improve conditions that are suitable for native
Restoration plantings. The most am- financial benefits, such as woodlots (arrow 5). In fauna than plantations of exotic species.
bitious goal is to attempt to reestablish the subsequent rotations, this balance might change over Such trials involving monoculture planta-
original forest ecosystem. Although the time (moving to arrow 6 and later to arrow 7 by using a tions of high-value native timber species
rules of ecosystem assembly are still greater variety of species). There may be greater scope for are now under way in many countries
debated (14), there are some empirical achieving multiple objectives by using several of these (SOM Text). Another approach is to use
options at different locations within the landscape mosaic.
data from several sites showing promising mixtures of species rather than simply
results (15–17). Two broad approaches monocultures (21). Recent experiments in
have been tested. Each involves a contrasting usually at high densities (92500 trees per ha), Costa Rica (22, 23) on commercially attractive
assembly rule, but each appears to offer and competitive interactions determine the native tree species suggest that mixtures in the
promising results under appropriate conditions. final forest composition (19). Species unable dry and humid tropics have the potential to
One approach is to use a small number of to tolerate open planting can be added once offer a number of benefits over monocultures,
fast-growing but short-lived tree species (i.e., canopy closure has occurred, either as seed- including production gains and reduced insect
equivalent to early successional pioneer spe- lings or by direct seeding. Such an approach damage. Another advantage of mixtures is that
cies) to create a canopy cover. These shade out was used in a forest restoration program they might provide small landowners a form of
grasses and weeds, diminish the fire hazard, after bauxite mining in central Amazonia insurance to protect them from uncertain future
and facilitate colonization of the site by a (SOM Text). In this case, over 160 species markets. However, the design of these mix-
wider range of species from nearby intact for- representing a range of life forms and succes- tures poses several key dilemmas. One ques-
est (1, 16). Under certain circumstances, these sional stages were planted after mining ceased tion is how many species are needed to
cover or nurse trees might be established by and topsoil replaced (Fig. 3). After 13 years, maximize these benefits; that is, what is the
direct seeding (18). The success of this low- the new forest, enriched by colonization by nature of the relationship between increased
diversity planting technique depends on the species from nearby intact forests, resembled diversity and any functional benefits?

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 REVIEW

Table 1. A summary of some of the different forms of reforestation that might be used when secondary forests are present or when some form of
planting is needed. Any combination of these techniques could be used in degraded landscapes depending on ecological circumstances and on the goals of
the land managers.
Plantings and plantations
Natural secondary forests
To restore biodiversity To supply goods and ecological services
Protect and manage Restoration plantings using small Tree plantation monoculture of exotic
natural regrowth: number of short-lived nurse trees: species:
Potentially able to supply Acquisition of further diversity dependent An efficient method of timber or food
a variety of goods and on colonization from nearby forest production for (mainly) industrial
services depending on remnants. Primary benefit is ecological users; in most circumstances it is less
the age and condition of services although can supply some successful in supplying many services.
the forest. goods depending on species present.

Protect and manage natural regrowth Restoration plantings using large number Tree plantation monoculture of native
plus enrichment with key species: of species from later successional stages: species:
Enrichment with Higher initial diversity that will also Useful to supply timbers of higher
commercially, socially, be supplemented by colonization commercial value and other goods
or ecologically useful from nearby forest remnants. such as fruits, nuts etc.; the longer
species can improve the Primary benefit is ecological rotations normally used may facilitate
value of these forests to services although can supply some an improved supply of ecological

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local communities or goods depending on species used. services such as watershed protection.
industry.
Direct seeding: Tree plantation used as a nurse crop with
The number of species that can be underplantings of native species not
established by direct seeding is otherwise able to establish at the site:
limited by seed supply but the An initial fast-growing nurse crop
establishment cost can be lower. supplying commercially useful timbers
Direct seeding can be used to or other goods can facilitate (e.g., via
initiate reforestation in open fields nitrogen fixation and microclimate
under appropriate conditions but it alterations) the subsequent
may be most useful when used to establishment of more species-rich
enhance diversity once some tree forests that supply a wider range of
cover is already present. goods and services.

Tree plantation mixtures of native

species:
Mixed-species plantations can,
potentially, supply a wider range of
goods and services than monocultures.
Biodiversity gains are greater than in
plantation monocultures but are mostly
still modest (usually less than five
planted species).

Although there is increasing evidence that (i.e., able to avoid one species outcompeting diversity of income streams available to the
functions such as production, litter decay, and and excluding others). It is clear that the nature landowner.
nutrient cycling can be enhanced as diversity of the diversity-function relation depends on
increases (24), there is also increasing accept- the particular species used and that the Making Reforestation Attractive
ance that simple taxonomic differences are an sampling effect can alter this relation (25). to Rural Communities
incomplete measure of diversity. Responses These theoretical problems are matched by The present scale of land degradation is such
may depend more on the diversity of function- some practical ones as well. Unless the market that it will only be overcome if large numbers
al groups (e.g., nitrogen fixers, slow- or fast- prices of the various species are similar, the of individual landowners or land managers
growing canopy trees, and bird-attracting overall value of the plantation will be reduced become involved in reforestation. But, large-
species) or on the diversity of functional re- as increasing numbers of lower value species scale tree planting or farm forestry is not
sponses within these groups rather than simply are added. commonly practiced by rural communities
on the number of tree species used in a This means there may be a limit on the use living in degraded landscapes, even though
plantation (8). But our incomplete knowledge of plantations to foster biodiversity at a par- many might practice some form of agro-
of tropical forest flora means it is rarely ticular site, although such plantations often forestry. This may be because most of their
possible to confidently categorize many spe- catalyze successional development in the land is needed for food production, but it is
cies into various functional groups apart from plantation understory (26). Trying to strike also because many rural people still have in-
relatively simple categories. Nor, for the same compromises between conservation and eco- secure land and tree tenure and are unwilling
reason, is it yet possible to define species nomically valued production may end up to invest in an activity from which they may
within a particular functional group that might generating suboptimal outcomes for each. derive little benefit. Reforestation can also be
have different functional responses. Nevertheless, mixtures may be useful if they unattractive because the initial costs can be
A second dilemma facing those wishing to are likely to enhance both ecological resilience high, whereas the direct financial benefits are
use mixed species plantations is that of and financial resilience of these new systems. delayed in comparison with a variety of
identifying species able to form stable mixes The latter would come from the greater other possible land uses the landowner might

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Fig. 3. (A) An aerial view of the open-cut bauxite mine at Trombetas in central
Amazonia that is located in a relatively undisturbed area of evergreen equatorial moist
forest. A reforestation program treats about 100 ha of mined land per year by using
stockpiled topsoil and by planting a variety of native species with direct seeding,
stumped saplings, or potted seedlings. (B) Within 10 years of establishment, most sites
have many more tree and shrub species than the number initially planted because of
seed stored in the topsoil or colonization from the surrounding forest. These new
species would have been brought to the site by birds, bats, and terrestrial mammals,
and most were species with small seeds. Overall it seems the reforestation program
has been successful in facilitating the reestablishment of both plants and animals to
the site, although more time will be needed for the composition and structure to
closely resemble nearby intact forest (SOM Text).

adopt (i.e., reforestation can have high op- than one species because they are interested in costs. This is particularly the case in land-
portunity costs). Even when tree planting is producing a variety of goods (13). scapes containing many smallholders (29).
undertaken, most landowners have often A third way to make reforestation attractive This means that some services (e.g., carbon
found it easier to use fast-growing exotic is to develop silvicultural systems by which sequestration) might be most easily provided
tree species than native species, about which plantations can be underplanted with crops that by large industrial plantations rather than by
there is much less ecological or silvicultural mature more quickly than trees, building on many small farmers. Such a market might then
knowledge. traditional and modern knowledge of agro- displace smaller farmers and thus generate
There are several ways by which re- forestry systems. These might be shade-tolerant significant social costs.
forestation might be made more attractive to agricultural cash crops (e.g., coffee, cocoa, and
landowners. One is to develop appropriate in- cardamom) or nontimber forest products such Forest Landscape Restoration
stitutional, legal, and policy settings (e.g., as rattans or medicinal plants (28). Again, there Most degraded tropical landscapes are a
providing secure land tenure, elimination of is often a significant local market for these mosaic of land uses and may include patches
‘‘perverse’’ incentives that favor deforestation species as supplies previously obtained from of intact residual forest and productive agri-
and forest degradation, and facilitating market- natural forests decline. cultural lands as well as degraded lands. It is
ing of forest goods) and to provide financial Lastly, reforestation might be more attract- rarely possible to reforest the whole landscape,
loans or inducements to make reforestation ive to landowners if they are paid for the eco- especially if it is also occupied by many small
attractive. logical services provided to those who benefit farms. Under these circumstances, forest res-
Another is to provide more information and from reforestation but who share neither the toration is usually done by concentrating on
technical assistance to landowners or commu- costs nor risks. Examples of payments for eco- particular sites. These might be riparian areas,
nities about the species suitable for planting, logical services provided by plantations in- buffer zones around residual forest patches,
their silvicultural requirements, and their mar- clude water, carbon, and biodiversity (29). corridors between forest areas, eroding areas
ket values. In many cases, the market prices of Such payments could make reforestation quite on steep hills, etc. However, the effectiveness
timber from slower growing native species are an attractive land use. Although this market of conserving biodiversity and restoring key
significantly higher than those for fast-growing has undergone significant growth in the past ecological functions that operate at landscape
exotics, and these prices are increasing as sup- decade, fundamental relations between forest scales (e.g., stabilizing hillslopes and hydro-
plies from natural forests decline (27). As the composition/structure and their functional logical processes) depends on these separately
supplies of low-value timbers from large in- characteristics, i.e., their ‘‘yields’’ of ecological restored sites complementing others in the
dustrial plantations flood the international services, are still poorly understood. This con- landscape mosaic. Individual decisions made
markets, the market niche for these high- tributes to the uncertainty of the market value by many small landholders are unlikely to
quality timbers may be a safer and more of these services. Further, the legal frameworks achieve this optimal outcome. This then prompts
valuable target for smallholders. Experience to allow trading are yet to be established in questions such as which parts of the landscape
to date suggests that smallholders who plant most tropical countries, and many of these should be reforested first, what type of re-
native tree species often prefer to use more markets are likely to have high transaction forestation should be carried out in particular

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(1988).
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landscape. Not only will these stakeholders grate agriculture and tree growing. One is the (1999).
differ in the extent to which they can or are drift of populations from country to cities, which 20. P. D. Erskine, Ecol. Manage. Restor. 3, 136 (2002).
21. A. C. Leopold, R. Andrus, A. Finkeldey, D. Knowles,
willing to share the costs and benefits of any appears to be widespread in many tropical For. Ecol. Manage. 142, 243 (2001).
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the size of their landholdings and in their in the markets for high-quality timbers and other Ecol. Manage. 175, 195 (2003).
23. D. Piotto, E. Viquez, F. Montagnini, M. Kanninen, For.
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and Ecosystem Functioning: Synthesis and Perspectives
at key resources such as water or soil con- services improve, there will be even more (Oxford Univ. Press, Oxford, 2002).
servation may be all that is possible. In the scope for making restoration and conservation 26. J. A. Parrotta, J. W. Turnbull, N. Jones, For. Ecol.
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27. T. K. Rudel et al., Global Environ. Change 15, 23
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relevant stakeholders in the planning and im- moving restoration from a site-based activity 28. M. S. Ashton et al., For. Ecol. Manage. 154, 431 (2001).
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(ITTO, Yokahama, Japan, 2004).
level that restoration can be used to comple- 30. W. M. Adams et al., Science 306, 1146 (2004).
Outlook ment the existing protected area network, 31. We thank D. Gilmour, M. Chapman, S. Brown, P. Dart,
The current rate of deforestation in tropical re- and it is at the landscape level that biodi- and colleagues within the Rainforest Cooperative
gions constitutes a major global biodiversity versity restoration and production (and hence Research Center, including N. Stork, C. Catterall, J.
Kanowski, J. Herbohn, S. Harrison, and G. Wardell-
crisis. This loss of biodiversity is significant, but poverty alleviation) can be most easily made Johnson, for their ongoing interest in tropical landscape
so too are the poverty levels of people who rely complementary. restoration and their constructive comments on the
on these forests and degraded lands for their ideas presented in this manuscript. We also thank D.
Kleine for creating the figures. Comments provided by
livelihoods. Both issues need to be addressed. References and Notes the anonymous referees helped improve our original
Conserving and actively managing the large 1. International Tropical Timber Organization (ITTO), manuscript. D.L. has an advisory role as the theme
areas of secondary forest that are now present Guidelines for the Restoration, Management and leader on ecosystem restoration for IUCN’s Commis-
Rehabilitation of Degraded and Secondary Tropical sion on Ecosystem Management.
is one way of doing this. Many rural com- Forests (ITTO, Yokohama, Japan, 2002).
munities are able to carry this out once ap- 2. C. R. Margules, R. L. Pressey, Nature 405, 243 (2000). Supporting Online Material
propriate government policies have been 3. T. M. Aide, J. K. Zimmerman, J. B. Pascarella, L. Rivera, www.sciencemag.org/cgi/content/full/310/5754/1628/
H. Marcano-Vega, Restor. Ecol. 8, 328 (2000). DC1
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