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 Author's personal copy
Opinion
Agroforestry: a refuge for tropical
biodiversity?
Shonil A. Bhagwat1, Katherine J. Willis1, H. John B. Birks2 and Robert J. Whittaker1
1
Biodiversity Research Group, Oxford University Centre for the Environment, University of Oxford, Oxford OX1 3QY, UK
2
Department of Biology, University of Bergen, Allegaten 41, N-5007 Bergen, Norway
As rates of deforestation continue to rise in many parts
of the tropics, the international conservation community
is faced with the challenge of finding approaches which
can reduce deforestation and provide rural livelihoods in
addition to conserving biodiversity. Much of modern-
day conservation is motivated by a desire to conserve
pristine nature in protected areas, while there is grow-
ing recognition of the long-term human involvement in
forest dynamics and of the importance of conservation
outside protected areas. Agroforestry intentional man-
agement of shade trees with agricultural crops has the
potential for providing habitats outside formally pro-
tected land, connecting nature reserves and alleviating
resource-use pressure on conservation areas. Here we
examine the role of agroforestry systems in maintaining
species diversity and conclude that these systems can
play an important role in biodiversity conservation in
human-dominated landscapes.
Biodiversity conservation in the tropics
In tropical regions, extensive conversion of forests and
agricultural intensification are typically identified as the
most prominent drivers of land-use change and biodiversity
loss [1,2]. Rates of deforestation in some parts of the tropics
are estimated to have increased since the beginning of the
21st century compared to those in the 1990s [3]. Moreover,
land-use pressure is compounded by the fact that some of the
worlds poorest people live in the tropics. It is estimated that
up to 86% of the population in some tropical countries lives
below the poverty line [4]. In addition, the population growth
rate in these regions is substantially higher than the popu-
lation growth rate of the world as a whole [5]. Therefore, any
approach that aims to mitigate tropical deforestation and
protect biodiversity should address the livelihoods and
needs of local communities.
Agroforestry intentional management of shade trees
with agricultural crops has emerged as one of the most
promising approaches to reducing deforestation in the
tropics while enhancing rural livelihoods [6]. Further-
more, it has been suggested that agroforestry systems can
alleviate the resource-use pressure on protected areas,
enhancing habitats for some wild species and increasing
the connectivity of landscape components, thereby mak-
ing conservation more effective [7]. However, the extent
to which agroforestry might provide a viable option for
biodiversity management outside formally protected
Corresponding author: Bhagwat, S.A. (shonil.bhagwat@gmail.com).
0169-5347/$ see front matter 2008 Elsevier Ltd. All rights reserved. doi:10.1016/j.tree.2008.01.005 Available online 24 March 2008
areas remains seriously under-researched (but see Ref.
[8]).
Here we review evidence from studies across the tropics
where species richness and composition of agroforestry
systems are compared with that of neighbouring forest
reserves. We treat this evidence from two perspectives.
First, we highlight the changing concept of pristine nature
by arguing that people have played a key role in shaping
many of the so-called pristine forests of today and empha-
sise that future approaches to conservation need to con-
sider agroforestry. Second, we recognise the so-called
matrix effect on species diversity in landscape mosaics
with native tree cover, and suggest that agroforestry sys-
tems can provide corridors that connect distant reserves.
Based on the evidence from the literature, we ask whether
agroforestry systems can offer a useful tool for biodiversity
conservation and conclude that they can play an important
role by providing habitat for many species in increasingly
human-dominated landscapes.
Conservation in human-dominated landscapes
Many landscapes that are considered pristine forests
today have, in fact, been under some form of cultivation
in the past [9]. Cultivation techniques included not only the
planting of trees but also soil-management strategies. For
example, evidence has recently emerged that the ancient
Maya manipulated and cultivated the landscape of the
Yalahau region in the northeast corner of the Yucatan
Peninsula in Mexico using algae from wetlands to enrich
upland garden plots, and cultivated trees within their
communities [10]. Similarly, there is substantial evidence
to indicate that the fertile Terra Preta soils were developed
by pre-Columbian native populations in central Amazonia
through the addition of large amounts of charred residues,
organic waste, excrement and bones [11]. A variety of
agroforestry crops are known to have been cultivated by
prehistoric populations in the tropics [12,13]. These in-
clude Brazil nut trees Bertholletia excelsa cultivated in
forest groves by the Kayapo Indians of the Brazilian Ama-
zon before European colonisation [14], the cultivation of
Boswellia papyrifera (the tree that yields frankincense)
over 2500 years ago in Africa [15] and banana cultivation in
the heart of the African rain forest at least 3000 years ago
[16]. The hunter-gatherer communities in Nkang,
southern Cameroon are also believed to have cultivated
bananas around 3000 years ago in addition to keeping
livestock [17]. Similarly, people inhabiting the lower mon-
tane rain forests of the highlands of Papua New Guinea are
261
 Author's personal copy
Opinion
believed to have maintained banana plantations around
6500 years ago [17].
There is a substantial lobby within modern-day conser-
vation that promotes protection of the largest possible
tracts of pristine landscapes [18]. Although large tracts
of forest habitat are necessary for conservation of, for
example, megafauna that have large home ranges, such
habitats are rarely as pristine as they are typically
represented [9]. In fact, it has long been known that people
have had a significant influence on landscape development
in tropical regions. There is ample archaeological evidence
of prehistoric human-induced environmental change, often
indicating degradation of natural landscapes [1921].
Recent work in applied palaeoecology is adding to a gra-
dual shift in perception that many of todays so-called
natural landscapes were, in fact, shaped by prehistoric
people [22]. Therefore, many areas conceptualised as pris-
tine forests might have regenerated from agricultural
fallows in the recent past. Long-term palaeoecological
records provide realistic estimates of the time it takes
for a forest to regenerate after abandonment from cultiva-
tion. For example, evidence of cultivation by humans
around 700 years ago has been reported from vast
stretches of so-called pristine rain forests in southern
Nigeria today [23]. Several other examples have shown
cycles of use, forest regeneration and re-use of land by
humans (see Ref. [24] and references therein). This work
emphasises the message that people are part of nature and
that the biodiversity found today is the result of past
human activities, as well as a combination of other eco-
logical and climatic processes [25]. A substantial body of
knowledge suggesting that todays pristine forests might
have been yesterdays agricultural fallows is, by necessity,
changing concepts and approaches to biodiversity conser-
vation [2528].
Figure 1. An example of a coffee plantation grown under the shade of native trees. In this coffee plantation in Kodagu, Western Ghats, India, a layer of coffee bushes is seen
in the understorey and a canopy of native trees is seen above this layer. Such plantations of shade-grown coffee in many tropical regions have canopy structure similar to
that of secondary forests. The shade of native trees provides habitat for many forest-dwelling species in otherwise highly human-dominated landscapes.
262
Trends in Ecology and Evolution Vol.23 No.5
Today, in some humid tropical landscapes, trees have
such a prominent place in farming systems that the differ-
ence between forests, old fallows and extensively managed
tree-crop plantations is not always apparent. Coffee plan-
tations in many tropical regions, for example, are grown
under shade of native tree species, making these planta-
tions resemble neighbouring forests in their structure [29]
(Figure 1). Such plantations provide habitat for many
species outside protected reserves in otherwise highly
human-dominated landscapes [7]. This role of agroforestry
plantations in biodiversity conservation is often over-
looked. Much emphasis is placed on fencing in forests
within strictly protected reserves and nature conservation
is largely regarded as being focused almost exclusively on
formally protected areas [30]. Although there is no doubt
that protected areas do contribute hugely to preserving
large parts of the biodiversity on Earth, such areas often
fail to cover the entire diversity of ecological habitats and
species and, as crucially, strict protection of such areas is
often resented by local people [31]. In a recent debate over
the future of tropical forest species, it has been suggested
that although some forest specialists need intact forest
habitat, the bulk of tropical species will be forced to persist
in degraded and secondary habitats outside forest reserves
in the future (see Ref. [32] and references therein). There-
fore, the need for biodiversity conservation in human-
dominated landscapes in the tropics is greater today than
ever.
Agroforest refuges for biodiversity
Recent research in tropical fragmented landscapes
suggests that small, isolated fragments, typically less than
100 ha in area (much of which is forest edge), are domi-
nated by common and invasive species and are poor in rare
and endemic species [33]. Agroforests are often very small
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Opinion Trends in Ecology and Evolution Vol.23 No.5

Figure 2. Countries represented in studies on species richness and composition of agroforestry systems. Twelve agroforestry systems and nine taxa in fourteen countries
across the tropics represent cases where a direct comparison was made between agroforestry systems and neighbouring forest reserves, and where species richness and
similarity in species composition were reported. For each country, A refers to the agroforestry system (As, allspice; Bn, banana; Bz, benzoin; Ca, cardamom; Cc, cocoa-
coffee; Cf, coffee; Co, cocoa; Ft, farm trees; Hg, home gardens; Jr, jungle rubber; Mf, mixed fruit orchard; Rt, rattan) and T refers to the plant or animal taxa (Br, birds; Bt,
bats; Hr, herptiles; In, insects; Mf, macrofungi; Mm, mammals; Pl, plants [lower]; Pn, plants [herbaceous]; Tr, trees).

in size, surrounded by open landscapes and resemble forest
fragments. It has therefore been argued that the species
distinctiveness (presence of rare or endemic species) of
such anthropogenic systems is frequently low, even though
their species richness (total number of species) might be
equal to or higher than that of neighbouring forests (e.g.
Ref. [34]). Noble and Dirzo [35], for example, suggest that
5080% of species from regional species pools typically
survive in agroforestry systems; however, many endemic
species are lost from agroforests [36]. Does this mean that
agroforestry systems have rather limited conservation
significance?
Although agroforestry systems might be impoverished
in richness of endemic or specialist forest species, and
distinctive in composition owing to their intensive man-
agement [35], the other side to this argument is that these
systems conserve biodiversity in habitat remnants in
otherwise open landscapes and provide enhanced potential
for species movements between habitat remnants. In a
literature search on the Web of Science database in Decem-
ber 2007 using keywords agroforestry and biodiversity,
we found 185 references. Of these, 36 made a direct com-
parison between agroforestry systems and neighbouring
forest reserves. In total, 12 types of agroforestry systems
and 9 taxa were represented in 14 tropical countries
(Figure 2). Many of these studies sampled multiple taxa
in forest and agroforestry landscapes, providing a total of
69 examples. Based on the published data, we calculated,
for each agroforest, the species richness as a percentage of
that found in the neighbouring forest reserve and
examined the similarity in species composition between
the two system types (summarised in Table 1; see
Appendix 1 in online Supplementary Material for details).
Although there is a wide variation across studies and
taxa, the mean values for richness in agroforestry systems
are greater than 60% of the forest values. Whereas com-
munities of bats and of lower plants (including bryophytes
and ferns) are richer (112139%) than in the neighbouring
forest reserves, the richness values for other taxa are lower
(6293%) (Table 1). The mean values of similarity in
species composition are between 25% (herbaceous plants)
and 65% (mammals). Although, in general, mobile taxa
such as bats, other mammals and birds display higher
similarity values and the values for plants are lower, in
certain studies agroforests show high levels of similarity to
neighbouring forest reserves (e.g. 91% for mammals, 98%
for insects and 100% for trees) (Table 1). The studies also
suggest that agroforestry systems are high in species
richness and more similar to neighbouring forest reserves
in species composition if (i) the forest land was fairly
recently converted to agroforestry plantation (e.g. Ref.
[37]); (ii) the management was less intensive (e.g. Ref.
[38]); and (iii) the canopy cover of native trees was high
(e.g. Ref. [39]). Furthermore, the studies reveal three major
reasons why agroforestry systems might be valuable for
biodiversity conservation.
First, many agroforestry systems are shown to be
important for the protection of species and habitats outside
formally protected areas. For example, Williams-Guillen
and colleagues [40] found that shade coffee plantations in
Nicaragua can serve as alternative wildlife habitats and as
corridors between forest fragments for mantled howling
monkeys Alouatta palliata. Similarly, research on the
distribution of birds on Khao Luang Mountain in southern
Thailand has shown that 3848% of bird species inhabiting
neighbouring forests are also found in mixed fruit orchards
[41]. Tylianakis and colleagues [42] also found a notable
overlap in the Hymenopteran communities of different
habitat types, including coffee agroforests and native forest
fragments in Ecuador, indicating that even intensively
managed land can make a valuable contribution to the
overall biodiversity of the landscape mosaic.

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Opinion Trends in Ecology and Evolution Vol.23 No.5

Table 1. Animal and plant taxa represented in the agroforestry systems studied and the richness and similarity in species
composition in relation to neighbouring forest reserves (see online Appendix 1 Supplementary Material for details)
Taxa reported Number of Agroforestry systems represented Richness compared to forest Similarity with forest Refs (in online
examples (abbreviated name [number of % (mean [range]) b % (mean [range]) c Supplementary
examples]) a Material)
Bats 3 Bn (1), Co (2) 139 (115186) 61 (5570) [11,15]
Birds 12 As (1), Bn (1), Cf (4), Co (3), Ft (1), Jr 92 (20179) 52 (1979) [3,4,11,15,18,20,23,27,
(1), Mf (1) 28,34,35]
Herptiles 1 As (1) 62 (62) 34 (1538) [18]
Insects 19 Bn (1), Co (10), Cc (1), Cf (6), Jr (1) 86 (44250) 49 (298) [1,5,810,14,26,2832]
Macrofungi 1 Cf (1) 89 (89) 61 (61) [4]
Mammals 3 As (1), Bn (1), Co (1) 93 (67121) 65 (4591) [14,18]
(excluding bats)
Plants (lower) 5 Co (2), Jr (3) 112 (77144) 42 (681) [2,3,30]
Plants 5 Co (2), Cc (1), Cf (1), Hg (1) 64 (25100) 25 (254) [6,22,28,30,36]
(herbaceous)
Trees 20 Bn (2), Bz (2), Ca (1), Co (6), Cc (1), Cf 64 (8213) 39 (5100) [3,4,68,1216,19
(4), Ft (1), Hg (1), Jr (1), Rt (1) 21,24,25,30,33]
a
Agroforestry systems: As, allspice; Bn, banana; Bz, benzoin; Ca, cardamom; Cc, cocoa-coffee; Cf, coffee; Co, cocoa; Ft, farm trees; Hg, home gardens; Jr, jungle rubber; Mf,
mixed fruit orchard; Rt, rattan.
b
Richness: values based on published data reporting species richness in agroforestry systems and neighbouring forest reserves. NB Values greater than 100 suggest
agroforestry system with species richness higher than that of neighbouring forest reserve owing to the presence of non-forest species.
c
Similarity: values based on published data reporting similarity in species composition between agroforestry systems and neighbouring forests. NB Differences in methods
involved between studies mean that the numerical comparisons shown here should be seen as only a rough guide.

Second, agroforestry systems maintain heterogeneity at
the habitat and landscape scales. For example, Parikesit
and colleagues [43] identified 12 different plant assem-
blages in Kebon Tatangkalan agroforests of the Upper
Citarum Watershed, West Java, Indonesia. These agrofor-
ests are maintained by small households spread across the
watershed and contribute to the highly heterogeneous
species assemblages in the regional flora. Kindt and col-
leagues [44] carried out complete tree inventories on 201
farms from four villages in western Kenya. They found
significant differences in tree species composition between
farms. The choice of shade trees left on their plantations by
individual farmers contributed to the overall heterogeneity
of trees in this agroforestry landscape. A study on Mount
Kilimanjaro has also shown that traditional coffee-banana
plantations, locally called Chagga homegardens, maintain
a multilayered vegetation structure similar to tropical
montane forests [45].
Third, trees in agroforestry landscapes reduce pressure
on formally protected forest reserves. Murniati and col-
leagues [46] studied the availability of forest resources in
mixed gardens and village forests in the northern part of
the Kerinci Seblat National Park, Sumatra, Indonesia.
They found that households that owned mixed gardens
depended much less on national park resources than
households which cultivated wetland rice fields alone.
Similarly, Masozera and Alavalapati [47] found that com-
munities which earned a high income from diversified
agricultural lands were much less dependent on forest
resources from the Nyungwe Forest Reserve in Rwanda
compared to communities which earned a low income from
agriculture. Research on trees outside forests in several
small-island developing states of the Pacific Ocean
suggests that the presence of such trees is valuable for
minimising deforestation, forest degradation and loss of
biodiversity from reserves [48].
Although networks of protected areas in the tropics
might currently provide habitat for many rare and endemic
species that prefer old-growth forest, it has been acknowl-
edged that these networks are insufficient to protect all
tropical biodiversity [49]. This is where agroforestry
systems can play an important role. Although agroforestry
systems cannot stand alone as conservation areas, they can
buffer existing reserves and provide corridors for persist-
ence and movement of species across landscapes (e.g. Ref.
[50]). Such systems offer a useful means for combating
species loss as a result of tropical forest fragmentation.
Countryside biogeography
For the last four decades, many approaches to conservation
have been rooted in MacArthur and Wilsons equilibrium
theory of island biogeography [51]. This theory set out to
account for the long-established relationship between the
number of species and (island) area one of the most
widely described general properties of ecological systems
[52]. Consistent with this theory, a loss in the size of
habitat is known to result in a proportional loss of species
[53]. Conversely, with an increase in the size of available
habitat, there will be a proportional increase in species
number. Thus, an agricultural landscape with high tree
cover should support more forest species than a landscape
without trees. The application of island theory led con-
servationists to view nature reserves as islands in a sea of
ruined habitat, and to focus on what configuration of
fragments provided the best return with regard to species
richness [54]. Partly as a result of this focus on island
theory, conservation biologists became so focused on
reserves that they have paid insufficient attention to the
significance of the properties of the landscape matrix
surrounding reserves [55].
Recent calls have been made, under the headings var-
iously of countryside biogeography [56], reconciliation
ecology [57] and conservation biogeography [58], for
research to establish how to create wildlife-friendly coun-
tryside landscapes. One part of this research agenda
should focus on the role of agroforests of various types,
to establish how effective they are as habitats and corridors
for wildlife within mixed countryside. Many of the worlds
agroforests provide a habitat-rich landscape matrix where
native species can survive outside nature reserves (e.g. Ref.

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[59]). Given that conservation decisions are often driven
more by political and economic rather than ecological
criteria and that the governance of nature reserves suffers
from high levels of corruption in many tropical countries,
conservation within reserves is constantly under threat
(e.g. Ref. [60]). Although protected reserves do play crucial
roles for conservation of biodiversity, a factor that will
contribute significantly to their success in the future is
the quality of the landscape matrix around reserves.
Finally, future climate change might put pressure on
species of conservation concern to migrate in response [61].
A landscape matrix surrounding reserves can provide
corridors through which species can migrate from one
reserve to another: a further reason why constituting wild-
life-friendly landscapes makes good sense. Therefore, agro-
forestry systems have the potential not only to provide
suitable habitat but also to provide corridors for the
migration of many forest-dwelling species that will face
the effects of changing climate in the future.
Conclusions
Long-term ecological knowledge that todays pristine for-
ests might have been yesterdays agricultural fallows
should be applied to the concept of naturalness of land-
scapes and approaches to biodiversity conservation today.
The ecological histories of many tropical landscapes cur-
rently within protected areas suggest that they were
impacted by human activities in the recent past. Although
the current protected area networks provide an important
conservation apparatus, further expansion of this network
might be limited by competing demands for other land
uses. Therefore, conservationists need also to look beyond
reserves and consider the importance of the surrounding
landscape matrix.
Agroforestry provides a potentially valuable conserva-
tion tool that can be useful for reducing land-use pressure
and enhancing rural livelihoods in tropical countries (Box 1).
The availability of useful tree species and other non-timber
forest products in agroforests can alleviate resource-use
pressure on conservation areas. Several examples across
the tropics have shown that a substantial proportion of
biodiversity of forest reserves is represented in agroforests.
The ability of many tree-covered but intensively used land-
scapes to support native species suggests that maintaining
and creating habitats in human-dominated landscapes can
help conserve a large proportion of biodiversity.
The literature reviewed suggests that agroforests with
less intensive management and high canopy cover have
high species richness and are more similar to neighbouring
forest reserves than intensively managed agroforests with
open canopies. Effective management of agroforestry sys-
tems, therefore, entails several aspects (see Box 1). First,
we suggest an integrated approach to reserve networks
that includes agroforestry systems as an important con-
servation tool. Second, we recommend that farmers who
maintain agroforests might, in particular circumstances,
be offered compensation in the form of various incentives
for biodiversity-friendly farming practices. Third, we advo-
cate that awareness and education programmes are crucial
to make farmers self-sufficient and to harness the potential
of agroforests for biodiversity conservation.
Trends in Ecology and Evolution Vol.23 No.5
Box 1. Management of agroforestry systems for
biodiversity conservation
It has been recognised that in order for the potential of agroforestry
to be harnessed effectively, farmers need clear incentives to plant
and protect trees [6]. Here we consolidate several recommendations
that might make agroforestry ecologically and economically viable
in the long term.
Landscape-based management
Although it is essential to maintain and cultivate shade-tree species
that are native to the region in question (instead of monospecific
shade), the variety of species cultivated is also of key importance
[62]. Similarly, keeping remnants of natural forest within agrofor-
estry plantations and redesigning annual croplands to include
features such as hedgerows can contribute to landscape-level
connectivity [63]. To sustain the conservation potential of agrofor-
estry systems, a balance between production and protection of
natural features in the landscape is necessary.
Incentives for conservation
It is important to recognise that farmers often have to forego part of
their income by maintaining forests on farmland. Although it is
essential that regional land-use planning is sensitive to the
economic needs of farmers [43], several incentives can be offered
to repay the cost of maintaining native trees on plantations [64]. Tax
relief for planting native shade trees, payment for environmental
services provided by agroforestry farms and soft credits for
ecologically beneficial farming practices have been suggested as
ways to make conservation lucrative to farmers [65]. A more direct
way of encouraging conservation in agroforestry systems is to
promote certification of shade-grown, biodiversity-friendly and
organic produce and to provide a secure market for certified
produce through farmers cooperatives [66]. Rewards for conserva-
tion are more likely to work than severe restrictions on agricultural
land use.
Education and training
In many tropical countries the levels of literacy are low. As a result,
communication of new government policies to farmers can be
challenging, and encouraging farmers to put them into practice
even more difficult [67]. Although national and regional pro-
grammes for planting native trees outside forests are essential
[48], community-based training programmes on techniques for
diversifying agroforestry plantations would also be beneficial to
farmers [68]. Furthermore, it would be crucial to educate farmers in
self-sufficiency to reduce their dependence on adjacent protected
areas. Self-sufficiency can be encouraged by providing training in
management and marketing of agroforestry produce [69]. Although
education in useful techniques and skills can enhance the benefits to
farmers from their agroforests, it can also alleviate some pressure
from reserves, thereby making protected areas more effective.
Acknowledgements
We are grateful to H.H. Birks, C. Garcia, P.K.R. Nair, K. Triantis, W.F.
Laurance and two anonymous referees for comments and helpful
suggestions on this manuscript. S.A.B.s current work at the Oxford
University Centre for the Environment is funded by the Leverhulme
Trust and a William L. Brown Research Fellowship of the Missouri
Botanical Garden. Collaboration with H.J.B.B. was initiated while he
held a Visiting Senior Research Fellowship at Jesus College, Oxford.
Supplementary data
Supplementary data associated with this article can
be found, in the online version, at doi:10.1016/j.tree.
2008.01.005.
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