Agriculture, Ecosystems and Environment 205 (2015) 25–35

Contents lists available at ScienceDirect

Agriculture, Ecosystems and Environment

journal homepage: www.elsevier.com/locate/agee

Review

Advances in knowledge of processes in soil–tree–crop interactions in

parkland systems in the West African Sahel: A review
J. Bayala a, * , J. Sanou b , Z. Teklehaimanot c, S.J. Ouedraogo d , A. Kalinganire a , R. Coe e,
M.van Noordwijk f
a
World Agroforestry Centre (ICRAF), ICRAF-WCA/Sahel Node, BP E5118 Bamako, Mali
b
Institut de l’Environnement et de Recherches Agricoles, Département Productions Forestières, 03 BP 7047 Ouagadougou, Burkina Faso
c
School of Environment, Natural Resources and Geography, Bangor University, Bangor, Gwynedd LLL57 2UW, UK
d
Institut du Sahel (INSAH), BP 1530 Bamako, Mali
e
World Agroforestry Centre (ICRAF), PO Box 30677, 00100 Nairobi, Kenya
f
World Agroforestry Centre (ICRAF), Southeast Asia Regional Programme, JL, CIFOR, Situ Gede, Sindang Barang, Bogor 16115, PO Box 161, Bogor 16001,
Indonesia

A R T I C L E I N F O A B S T R A C T

Article history: The ‘parklands’ that form the most widespread farming systems in the Sahelian zone of West Africa are
Received 6 August 2014 farming systems in which annual crops are grown under scattered trees preserved from the natural
Received in revised form 24 February 2015 vegetation by farmers following clearing the woodlands to make crop fields. Being mixed agricultural
Accepted 28 February 2015
systems, the interactions between trees and crops have always been a key element determining the
Available online 11 March 2015
management options applied by farmers. This has also attracted the attention of scientists, with the first
studies of this nature dating back to the 1960’s. A combination of field trials, observational studies and
Keywords:
modeling has been deployed to understand soil–tree–crop interactions, including sharing of growth
Ecosystem services
Management
resources by the system components. Based on this understanding, management options have been
Modeling discussed with farmers. The most common experimental designs used are transects from the base of tree
Productivity trunk toward the open field, concentric zones around trees or sectors under trees where various
Sub-Saharan Africa experimental treatments are applied, depending on the objectives of the studies. Key findings
Sustainability disentangled the contributions of each component of the system to the island of high fertility around
trees using isotopic techniques, tree effects on adjacent C4 cultivated plants and alternative C3 crops that
tolerate tree shade. Tree management practices such as pruning have been tested to modify the crown
architecture and the patterns of root distribution in order to improve light availability and resource use
efficiency. Increases in land use intensity require more active management by farmers of tree
regeneration. Despite scientific advances, there are still some methodological challenges in determining
the “park effect”, the tradeoffs and synergies between and among goods and services, and how to boost
the provisioning, supporting and regulating functions of such agroforestry systems. Providing such
ecosystem service functions is critical in the quest for ensuring food security while achieving adaptation
and mitigation goals in vulnerable environments like the drylands.
ã 2015 Elsevier B.V. All rights reserved.

Contents

1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
2. Complexity of the systems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
2.1. Multiple interacting components . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
2.2. Species diversity and diverse styles of management . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
2.3. Diverse spatial and temporal scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
3. Research approaches and methodologies employed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
4. Plot scale processes in soil–tree–crop interactions in parklands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29

* Corresponding author: Tel.: +223 20 70 92 20/77 71 41 90; fax: +223 44 90 18 07.

E-mail address: j.bayala@cgiar.org (J. Bayala).

http://dx.doi.org/10.1016/j.agee.2015.02.018
0167-8809/ ã 2015 Elsevier B.V. All rights reserved.
26 J. Bayala et al. / Agriculture, Ecosystems and Environment 205 (2015) 25–35

5. Current understanding of soil–tree–crop interactions in parklands ... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31

6. Conclusion and perspectives . . . . . . . . . ..................... .... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
Acknowledgements . . . . . . . . . . . . . . . . ..................... .... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
References . . . . . . . . . . . . . . . . . . . . . . . ..................... .... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33

1. Introduction
Although parklands occur in other areas of the world, such as
Southern Africa and the Mediterranean regions, it is in semi-arid
West Africa that these agroforestry systems are most widespread
(Kessler, 1992; Pullan, 1974 Zomer et al., 2009). Parkland is a
traditional land-use system that involves retention and/or
introduction of trees or other woody perennials in agricultural
fields and managing them in combination with crops and livestock
(Boffa, 1999; Bonkoungou et al., 1997). The aim is to benefit from
the positive ecological and economic interactions that take place
between the components. Parklands are generally made up of
several tree or shrub species that constitute an important source of
medicine, food, fodder and fuelwood as well as cash to local
communities (Bayala, 2002; Gijsbers et al., 1994; Jonsson et al.,
1999; Lamien et al., 1996). They also play a major role in preserving
the resources through improving soils (Bayala et al., 2006;
Takimoto et al., 2008). Despite their important socio-ecological
role, parkland systems have been reported to be degrading due to
increasing aridity (Gonzalez et al., 2012) and human pressure
hindering the natural regeneration of trees (Bayala et al., 2011;
Gijsbers et al., 1994; Maranz, 2009). Indeed, besides natural factors
(low and highly variable rainfall, long dry season of at least nine
months and periodic droughts that may last several years), much of
the Sahelian agriculture is locked in a spiral of degradation as a
result of growing demographic pressures, rapid social changes,
new policy environments, and cash-crop commercial farming
systems by encouraging the removal of unplanted trees (Ræbild
et al., 2012). Symptoms of parkland degradation include the
reduction in tree density and the absence of regeneration
(Augusseau et al., 2006; Gonzalez, 2001; Gonzalez et al., 2012;
Maranz, 2009; Wezel and Lykke, 2006). This has led, in many areas,
to lower the system’s buffer capacity as a result of the loss of the
tree component from the system, which is essential for the proper
ecological functioning via regulating soil temperature, soil
humidity, nutrient storage and cycling, as well as the dependent
soil biological processes that determine soil quality (Bayala et al.,
2014; Kater et al., 1992; van Noordwijk et al., 2014). Rehabilitation
of the degrading parkland systems should be grounded on a sound
understanding of the ecological functioning of the systems. Due to
the considerable potential benefits of complementarity in terms of
resources use between agroforestry system components, some
quantitative attempts have been made to analyze the productivity
of the systems in terms of resource capture and use by the
components (Bayala, 2002; Bazié et al., 2012). Indeed, a
fundamental understanding of how agroforestry system compo-
nents utilize available resources is crucial for determining species
combinations, planting arrangements, tree spatial densities and
management strategies suitable for different locations and
different farmer objectives and for the long-term sustainability
of the system (Ong et al., 1996). Research carried out to understand
how better trees and annual crops can be combined dates back to
the 1960’s starting with Faidherbia albida (Boffa, 1999; Charreau
and Vidal, 1965). This interest is due to observations of better crop
performance under this species in link with its reverse phenology
losing leaves during the cropping season and therefore competing
less or not at all with the associated crops (Roupsard et al., 1999).
Experimenting with such systems is not easy due to the
complexity, the diversity of species, the multiple scales at which
interactions occur and the long lifespan of trees. Hence research
has had to use a combination of experiments, observational studies
and models to understand processes and make predictions about
managing practices. This paper is aimed at providing an update
mainly from the 1990’s of what has been done in the area of soil–
tree–crop interaction studies in parklands of the Sahel and
suggestions on the way forward.
2. Complexity of the systems
The complexity of parklands resides in the fact there are
multiple interacting components, diverse species composing each
component, various ways of managing these components, diverse
scales in space (important processes as micro, plot and landscape
scales are different) and time. These three together make it hard to
study processes, but also make it essential if we are to understand
parklands.
2.1. Multiple interacting components
Parklands in the Sahel are formed of selected trees from the
original woodland after clearing the natural vegetation for
cropping, though trees can also be planted and natural regenerated
individuals be protected. At the time of clearing woodland, farmers
remove most of the vegetation, leaving only selected trees. As a
result and despite the diverse mixture of species in natural
vegetation, parklands are dominated at field level by a few tree
species (Boffa, 1995; Ouédraogo, 1995) mostly selected for their
food, wood and medicinal functions (Faye et al., 2011). At
landscape level the diversity can be higher than what is observed
at field level. For instance, more than 60 woody species have been
recorded having fodder function in the parklands of Sahel
(Arbonnier, 2000; Food and Agriculture Organization (FAO),
1992; Hansen et al., 2008; Le Houerou, 1980; Sanon et al., 2007;
von Maydell, 1983). Indeed, trees contain higher N content
(16–33 g kg 1) during the dry season than grasses making them
the major source of protein to livestock (Breman and Kessler, 1995;
Le Houerou, 1980). Livestock also feed on crop residues and fodder
tree species on the farmed fields and by so doing they deposit
manure in the fields, but also remove crop residues, which can be
important for protecting soil structure and retaining moisture. The
woody component, comprising trees and shrubs, is mixed with
cultivated staple food crops dominated by cereals (millet, sorghum
and maize). The cereals are generally intercropped with legumes
like cowpea or rotated with groundnut for soil restoration. Besides
these main crops, cash crops like cotton and sesame are also
cultivated.
Selected trees in parklands are spatially distributed in three
zones: house fields or compound/homestead fields, village fields
and bush/fallow fields (Bayala et al., 2011; Prudencio, 1983). This
spatial arrangement starts from homestead and progressively
moves outward toward the village/crop fields and bush/fallow
fields successively (Bayala et al., 2000; Ouédraogo, 1995; Raison,
1988). The management differs in relation with the various field
types. Homestead fields are more planted with exotic fruit trees
like mangoes and citrus and received more manure and wastes
from the households. They are also cropped for longer periods
(Bayala et al., 2011; Prudencio, 1983). Less fertilizer is applied and
 J. Bayala et al. / Agriculture, Ecosystems and Environment 205 (2015) 25–35
farming period is shorter moving from the homestead to the bush
fields.
2.2. Species diversity and diverse styles of management
The dominant tree species of parkland systems are Adansonia
digitata, Parkia biglobosa, Vitellaria paradoxa, and F. albida; the
latter being appreciated by local populations as a good parkland
species because of better crop production underneath due its
reverse phenology and other favorable traits to the associated
crops (Roupsard et al., 1999). Besides the parklands of F. albida, P.
biglobosa and V. paradoxa, other existing ones are as those of
Balanites aegyptiaca, Borassus spp., Lannea spp., Prosopis africana,
Sclerocarya birrea, Sterculia setigera, named after the dominant
species. A few of the parklands are almost (more than 90% of the
individuals) mono-specific (F. albida, Borassus spp., etc.) whereas in
others some species dominate but are mixed with a large range of
species making the system very complex to study for the
interactions with agricultural crops and livestock. The livestock
component is even more difficult because of its mobility, leading to
a dearth of studies of its interactions with trees. Therefore this
aspect will not be the focus of this review, but we hypothesize that
disentangling this topic could make a significant contribution to
this field of research because of its potential impact on vegetation
dynamics and nutrient flows. Shrubs are repeatedly coppiced at
the onset of the rainy season to prepare the fields for sowing and at
each cropping operation throughout the rainy season. Fodder trees
are pruned during the dry season to feed the livestock during the
period of forage shortage whereas some fruit tree like P. biglobosa
and V. paradoxa are pruned when old to rejuvenate them for better
fruit production (Bayala et al., 2008b). Finally other species like A.
digitata, Bombax costatum, Lannea microcarpa, etc., are pruned on a
yearly basis to harvest their fruit or leaves.
The cultivated crops in parklands can be categorized into two
types which are C3 and C4 plant types of photosynthesis pathway
and that have some implications when associated with shade trees
(Fig. 1). One advantage of the C4 pathway of photosynthesis is the
absence of the photorespiration process making them more
adapted to full sun environment. In turn they are less adapted
to shade compared to C3 which are able to modify the morphology
and anatomy of their leaves to allow them to harvest more light
energy in shade conditions. Therefore, the net photosynthesis of C3
plants (Gossypium herbaceum, Manihot esculenta, Colocasia
Fig. 1. Higher light saturation point (1) and lower compensation point in C4 (2) than
in C3 plants (reproduced from http://www.marietta.edu/spilatrs/biol103/photo-
laB/c3v4expl.html).
27
esculenta, etc.) is less reduced under shade conditions compared
to C4 plants (Zea mays, Sorghum bicolor, Pennisetum glaucum, etc.)
(Hay and Porter, 2006; Osborne et al., 2008,b; Sanou et al., 2012a,
b). C4 pathway plants have generally a higher water use efficiency
(WUE) compared to C3 plants when placed in the same
environmental conditions (Ripley et al., 2007). C4 plants are
similar or more sensitive to water stress compared to C3 pathway
plants despite their higher WUE (Ghannoum, 2009). The reactions
of the two crop types to nutrient are variable (Ripley et al., 2008).
Despite the above general characteristics, it is still not clear which
one of the factors is the most limiting in a given parkland system
and under given climatic conditions unless evaluated in real
situation.
In addition to tree diversity, agroforestry systems also help to
conserve plant, animal and microbial biodiversity by offering
diverse niches and food both aboveground and belowground. In
the belowground biodiversity conservation domain, there is dearth
of data (Diedhiou et al., 2009; Gnankambary et al., 2008; INERA,
2000).
2.3. Diverse spatial and temporal scales
There are several imbricated space scales going from the
individual to the plot, the watershed, and landscape levels. As
stated above, there is also a spatial organization of the parklands
around homesteads with different types which are compound,
village and bush fields (Bayala et al., 2011; Prudencio, 1983). Finally,
lowland ecology is different from that of upper lands and support
different tree species and crop types.
With regards to the time scales, it is important to highlight the
fact that some components are annual while others are perennial
with different life cycle lengths. For instance, short cycle crop
varieties have been commonly developed as a way to address the
issue of reducing rainy season length. Due to its possibility to
increase or better recycle nutrients, parkland of species like F.
albida are cropped for long time period compared to other species
(Boffa, 1999). Compound field is cropped longer time due to the
application of manure and wastes from the household (Prudencio,
1983).
3. Research approaches and methodologies employed
Research has always proceeded by simplification of systems,
and parkland systems did not escape from such an approach. The
first aspect of the simplification has led most of the investigators to
look at the individual effect of one tree species at a time. The
second area of simplifying the issue has been to consider grain
yield of associated crops as the only important property of the
system to assess. A third simplification has been to only consider
the effects at a single scale, typically the ‘plot’ consisting of a single
tree and adjacent cropland. A fourth simplification is to concen-
trate on processes happening at time scales from the day to the
year. And a fifth one is to decouple social and biophysical
investigations. The various methods used to study soil–tree–crop
interactions in the Sahel are described in the next sections, with
emphasis on the plot scale. This was meant to help focusing on
natural processes and therefore removing the need to pay so much
attention to the more complex human-mediated processes.
Appropriate methods are determined by research objectives.
The overall aim of much research on tree–crop interactions in
parkland systems is to predict the effect of changes, in particular
changes in tree density, species mix, or management on a number
of factors including but not restricted to soil, micro-climate and
associated crops. A common research approach proceeds by
observing that crop yield (or soil nutrients, or any other response of
interest) is greater under existing tree canopies than away from the
28 J. Bayala et al. / Agriculture, Ecosystems and Environment 205 (2015) 25–35
trees and thus concludes that increasing the tree density will
increase overall crop production. That is what has been done in
observational studies conducted on scattered trees without
considering their distribution pattern, mix and tree age which is
unknown. Three main approaches (Fig. 2) have been used for
observational studies of the interactions between components of
the parkland systems.
The first is the transect approach consisting of monitoring
quadrat plots laid out along a gradient running from the base of
tree trunk to the open area following the cardinal compass
directions (Kater et al., 1992; Kessler, 1992). A second modality of
the transect approach is to monitor crop performance between
pairs of trees (Boffa, 1999; Boffa et al., 2000; Charreau and Vidal,
1965). Boffa et al. (2000) have used rectangular blocks with trees of
the same species at each end. Transect approaches were found to
not properly deal with directional variations and high variability of
soil properties that can occur over very short distances in the Sahel
(Bayala et al., 2002; Louppe et al., 1996). Therefore, some authors
have opted for concentric zones around an individual tree going
away from the base of the tree trunk to the open area (Bayala et al.,
2002; Louppe et al., 1996). The concentric zoning approach aims to
reduce directional biases due to leaf and rain effects as well as
micro-variability in soil fertility (Boffa, 1999). Another modality of
the zoning approach was to subdivide the tree influence area,
considered as the vertical projection of the crown, assimilated to a
circle based on its average diameter in E–W and N–S directions, in
Fig. 2. Different experimental designs centered on individual trees used in studying
soil–tree–crop interactions in parkland systems in West Africa Sahel.
sectors delimited by the cardinal and ordinal directions from the
base of the tree trunk (Bazié et al., 2012; Kho et al., 2001; Nur
Osman et al., 2011). If zonation designs are more able to capture the
directional variability they fail to address the park effect as does
the sector method. In addition to the search for the most
appropriate design, the way data have been collected and analyzed
poses also some concerns. Most of the data did not exclude the area
occupied by the trunks of the trees and that may have led to some
underestimations of the beneficial effects of trees. Except for the
sector approach, most of the designs, although systematic, cannot
allow a real randomization of the gradient of tree effect. Yet most of
the data have been analyzed using ANOVA, while in such cases, it is
not possible to calculate a valid estimation of error as residuals of
zones or plots on transects are correlated (Sanou, 2010; Wilson
et al., 1998). While the variation of the data according to other
factors (year, species, etc.) could still be handled using ANOVA
(general linear model), the variation due to the tree gradient effect
may be analyzed better using pairwise t-tests (Sanou, 2010). The
above-mentioned methodological difficulties may have led to
some biases, but it is difficult to determine the direction or the
magnitude. A more systematic way of assessing the park effect is
the yield mapping method. It proceeds by dividing the plot of
parkland under investigation into small quadrats (2 m  2 m in
size), assessing crop yield (grain and biomass) within each quadrat
of 4 m2 and collecting all explanatory variables (soil properties) of
the crop yield within the same quadrat (Fig. 3; Ellis, 2013). One of
the already obvious issues is the cost associated with labor and soil
analysis of these extensive experiments. However such high cost
could be reduced by using geostatistical methods to more
efficiently investigate the defined area, once initial data describing
the spatial variability have been gathered. All the above studies are
observational and not experimental in that they measure what is
there but do not attempt to manipulate anything by imposing an
experimental treatment. Furthermore, changing the detail of how
studies are done around existing trees (concentric circles, trans-
ects, how many crown widths, etc.) does not change the basic
limitations of the approach as described above. While the
observations may be valid the conclusion is flawed. The problems
with such observational studies, however carefully done, are
numerous. For example, it is not possible to tell how far away from
the tree is not influenced by it, and hence can be taken to represent
‘without trees’. It is also not possible to tell whether the apparent
benefits of tree presence scale in a simple way. If, for example, the
high soil nutrients under a canopy are due to animals congregating
there then adding more trees will not increase the total added
nutrients, it will simply distribute the same pool across a larger
area. There are also subtle biases due to the fact that parklands are
managed systems. For example, it is highly likely that if a farmer
observes a tree having an apparent negative effect, this tree will be
cut down. Thus the trees remaining represent only ‘good’ ones.
Another example is the hypothesis – difficult to easily disprove at
plot level unless using isotopes – that existing trees occur in
preexisting fertility hotspots (Bayala et al., 2006). In turn, at
landscape level, Hadgu et al. (2009) have observed that areas with
higher tree densities corresponded to those of higher soil fertility
(available P and total N) and higher crop productivity in Tigray
region in Ethiopia.
While observational studies are appealing and easy to conduct,
it is also important to do experiments. An experiment to determine
the effect of adding trees does precisely that: compares areas that
are equivalent at the start of the study and into which trees are
planted in some and not in others, the allocation being made at
random. However this approach also has limitations. The most
obvious one is the time involved. There is evidence that beneficial
effects of F. albida on undersown crops take up to 20 years to
appear, and the timescale of other species may be longer. Yet few
 J. Bayala et al. / Agriculture, Ecosystems and Environment 205 (2015) 25–35
Fig. 3. Maize (Zea mays) yield mapping in Sokouraba in Burkina Faso where microsite of dead tree could be seen while the effects of living trees were not perceptible due to
small size of trees (source Ellis, 2013).
projects operate with that sort of time horizon – researchers and
donors want results faster. Other difficulties with design experi-
ments are the need to work with small plots, hence not being able
to study processes at larger scale, and the problem of ensuring that
plots are not interfering with each other (e.g. are no-tree plots
really not influenced by trees in neighboring plots?).
Some manipulative experiments are possible and useful. For
example, transfer experiments have been used, which consist of
transferring soil from under the tree to a corresponding plot in the
open area and vice-versa (Dembélé, 1994; Jonsson, 1995). This is an
attempt to separate out the effects of soil from those of microclimate.
A more drastic way of approaching the problem of presence/absence
of trees was applied by Dossa et al. (2012) who removed the woody
component to obtain treeless plots and compared these with plots
which still contained trees/shrubs. However, such an approach is still
missing out the different combinations of species and only allows
investigation of single factors. Furthermore, such method was
certainly not able to exclude the effects of decomposing biomass
formed of leaves and fine roots from the uprooted woody
component. Pruning of both roots (Bayala et al., 2013) and branches
(Bayala et al., 2002; Kessler, 1992; Tilander et al., 1995) and assessing
their effects on associated crops have also been realized. Due to
transient nature of the effects of pruning, there is a dearth of
information on its long term benefits. Alternative C3 crops that are
shade tolerant compared to cereals (C4) were also evaluated under
trees (Pouliot et al., 2012; Sanou, 2010; Sidibé, 2010). Finally, few
planted systems have been tested and reported in the literature. They
have used the strip approach with distance from tree rows (Ndiaye
et al., 2000; Tilander et al., 1995). In addition to field observations,
modeling has recently been used to identify the most limiting factors
when combining trees and annual crops (Bayala et al., 2008c;
Coulibaly et al., 2014; Kho, 2000; Mayus, 1998). In that line, yield
mapping coupled with modeling may help to find the solutions about
park effect, species mixture and tree density in “what-if” scenario
analysis.
The implication of these methodological constraints is that
conclusions and predictions about system response to changes
must be made on the basis of multiple sources of information.
Making progress requires combining observational studies (most
useful for identifying phenomena to explain and for measuring the
rates of processes), manipulative experiments (for testing specific
hypotheses) and models (for assembling information and making
predictions). When interpreting the results of observational
studies it is important to make sure that multiple explanations
of the observations have been examined and the conclusions are
sound.
29
4. Plot scale processes in soil–tree–crop interactions in
parklands
Despite the methodological difficulties, mixing C4 plants with
heavy shade-casting trees will certainly lead to yield reduction and
this is linked with the C4 plant photosynthetic pathway (Fig. 1).
This calls for management options to get the mixture correct or to
influence the interactions in such a way that the overall
performance of the system is improved and sustained. Even
though trees at farm scale can be considered to being a risk
reduction strategy (providing food during poor rainfall and
buffering harsh climate effects) rather than an overall produc-
tion-enhancer, their contribution to landscape scale issues for long
term production and in the context of changing climate is not yet
fully investigated. Thus the results of simulating different
management options under different climate scenario using
WaNuLCAS (Water, Nutrient, and Light Capture in Agroforestry
Systems) model revealed that the options of tree canopy pruning to
reduce shading while reducing fine root density was key to
parkland adaptation to a changing climate (Coulibaly et al., 2014).
In addition to microclimate modification, soil fertility improve-
ment seems be an accepted evidence (Fig. 4). However, controver-
sy still exists as to whether tree really contributes to soil fertility,
recycles it, or just “harvests” it from within its laterally extensive
root zone, and concentrates it (Bayala et al., 2006). Such question
still remains unanswered, even for species like F. albida, which is
leguminous. For instance, Maïga (1987) reported higher cereal
yields in control plots associated with F. albida. Such findings were
considered to support the indications that it is not F. albida tree that
fertilizes the soil but the high soil fertility that enabled its
establishment and proper functioning (Kessler, 1992). Such
conclusion does not consider the fact that F. albida have small
leaves that are easily transported and spread widely by the wind
even though we should not forget the contribution from root
turnover, microclimate increasing soil biota, etc. More homo-
geneously distributed leaves may have spread their effect on the
cropland for F. albida compared to those with larger and heavier
leaves that fall and stay in the immediate tree surroundings.
Indeed, while pruning yielded 15–40 kg 1 tree 1 year 1 of leaf
biomass (Depommier and Guerin, 1996), mean leaf litter was found
to be only 8 kg tree 1 using litter traps installed under F. albida
trees (Zomboudré, 2009). This illustrates very well the fact that
there are counter facts of the fertility preceding the trees. Isotopic
studies have helped to show trees’ higher contribution to soil
carbon building up in mixed perennial/annual agricultural
systems, compared to that of the crop component (Bayala et al.,
30 J. Bayala et al. / Agriculture, Ecosystems and Environment 205 (2015) 25–35
Fig. 4. Nutrient balance, growth resource sharing and environmental modification on associated crop development under trees in parklands in the Sahel.
2006; Jonsson, 1995). Despite such findings, it can still be argued
that, if the parkland effect on soil fertility is due to lateral
redistribution of nutrients by animals and/or roots or of pre-
existing higher soil fertility (Geiger et al., 1994), there are no gains
in productivity as a whole with a higher tree density (Kho et al.,
2001). But it can also be argued that the system is preserving
something which otherwise will be depleted. In addition, in the
Sahelian environments, cation exchange capacity (CEC) strongly
depends on the presence of organic carbon, which of course has an
impact on soil fertility. Trees have been reported to have a
significant impact on CEC in studies comparing under and away
tree plots with all their limitations. For instance, Samba (1997)
found a CEC 15–27% higher under Cordyla pinnata in comparison
with the treeless zone (CEC = 2.86 cmol kg 1). It is known that CEC
lower than 4 meq/100 g makes the use of mineral fertilizers non
economical, leading to the need to enhance this compensatory role
of trees in the Sahelian soils (Kater et al., 1992). Thus, trees, and
particularly agroforestry systems, are nowadays widely acknowl-
edged as valuable land management options.
A number of reports have highlighted the fact that upperstorey
(trees) and understorey (crops) components affect each other via
growth resources sharing (Fig. 4). The aboveground competition is
related to light, temperature, air relative humidity, and rain
interception (Jonsson et al., 1999; Kater et al., 1992; Kessler, 1992;
Sanou et al., 2012a). The belowground competition is about water
and nutrients use and it is generally expected that trees and crops
are occupying different soil layers (Schroth, 1995). However, field
data have shown that there is very often overlap of roots of both
plant types, thus exacerbating the belowground competition
between them (Bayala et al., 2004; Jones et al., 1998). Indeed,
whether competition or facilitation/complementar takes place
depends on the species and the ecological context. F. albida with
reverse phenology has been reported to stimulate crop yield,
particularly cereals as they require more light, being C4 plants.
Similarly, species with a canopy allowing some light to reach
underneath, even when in leaf (A. indica, Borassus spp., H. thebaica,
etc.), will still positively impact cereal production (Boffa, 1999;
Moussa, 1997; Yaméogo, 2008). When the leafing of both crop and
tree coincide for most of tree species, there is competition for light
for which C4 plants are more affected than the C3 crops (Sanou
et al., 2012a,b,b). That is why, under species like V. paradoxa and P.
biglobosa, crop performance may be significantly depressed (Kater
et al., 1992; Kessler, 1992) but there is also some evidence of crop
performance improvement depending on the species, density,
management, season characteristics, etc. (Boffa et al., 2000; Sabiitti
and Cobbina, 1992). As a general trend, a meta-analysis revealed
that yield increases were lower, relative to the control under
parkland trees, where annual average rainfall was higher than
800 mm and on sites where the yields of the control plots were
larger than 2 t ha 1 (Bayala et al., 2012). The lack of unambiguous
cereal yield benefits from the traditional practice of parkland trees
is probably realistic. Thus, testing shade-tolerant crops such as yam
(Dioscorea sp.), sweet potato (Ipomea batatas), taro (C. esculenta),
cocoyam (Xanthosoma sagittifolium) and cassava (M. esculenta) may
be a useful alternative and has proven to be beneficial to farmers
(Johnston and Onwueme, 1998; Onwueme and Johnston, 2000;
Pouliot et al., 2012,b; Sanou et al., 2012a,b; Sidibé, 2010). An
alternative to that will be to breed shade tolerant varieties of staple
food crops like cereals in the Sahel region. Recent remarkable
advances in genomics and bioinformatics in crop improvement
open up ways of breeding C4 plants that will be tolerant to shade
and that will associate well with the trees in agroforestry systems
 J. Bayala et al. / Agriculture, Ecosystems and Environment 205 (2015) 25–35
(Furbank et al., 2009; Gowik and Westhoff, 2011; Mochida and
Shinozaki, 2010). Pruning, as a tree management tool, has also been
successful for increasing cereal yields (Bayala et al., 2002; Bazié
et al., 2012). However, trees are traditionally maintained in
croplands for many purposes in addition to sustaining cereal
yields. A range of products is generated from the trees and
evaluations have shown that fruit production was enough to
compensate for the loss in cereal production (Bayala et al., 2008b;
Boffa, 1995). Given the above-evoked products and services, it may
be concluded that there is a need for a more comprehensive
analysis at household (farmers’ motivations to keep parklands) and
landscape (ecosystem services) scales of the multiple benefits and
services provided by parkland trees (Bayala et al., 2012). Indeed,
Bargués et al. (2014) and Bazié (2013) showed that preserving V.
paradoxa trees in parklands was vital to reduce runoff and increase
water infiltration revealing its ecophysiological/ecohydrological
role at landscape scale.
5. Current understanding of soil–tree–crop interactions in
parklands
The greatest opportunity in simultaneous agroforestry practi-
ces is the filling of niches within the landscape where resources are
currently under-utilized by monocrop agriculture (Ong and Leakey,
1999). However, microclimate modification by parkland trees (e.g.,
reducing supra-optimal temperatures) is not always about under-
utilized resources but may also be about increasing resource use
efficiency, for example by avoiding supra-optimal temperatures
(Sanou et al., 2012a,b,b; van Noordwijk et al., 2014). Mixing
different plant species is always associated with competition and
facilitation/complementarity as a result of niche differentiation
(Bruno et al., 2003; Kelty and Cameron, 1995). Potential successful
practices of mixing trees and crops are those in which competition
is out-weighted by facilitation ensuring higher and/or more stable
performance of the systems (Ong and Leakey, 1999). However,
scientific studies of the phenomena of facilitation and competition
in agroforestry in the Sahel are scarce because of the complexity of
tree–crop interactions (Bayala, 2002).
The entry point in these studies has been the assessment of the
yield of the associated crop followed by the search for explanatory
variables. There have been some studies showing that tree
influence zone is associated with a gradient of soil fertility
declining as one moves away from the tree base (Bayala et al.,
2006; Bayala and Ouedraogo, 2008; Breman and Kessler, 1995). As
most of the trees were not planted, the hypothesis that the island of
high soil fertility associated with the presence of trees reflects a
priori difference in fertility, allowing for better tree establishment
on richer sites, has been evoked by some authors (Bayala et al.,
2006; Brouwer et al., 1993; van Noordwijk and Ong, 1999).
However, there is now emerging data in the literature that shows
that the higher carbon under trees is due to their direct positive
impact on soil organic matter formation (Bayala et al., 2006;
Jonsson, 1995; Takimoto et al., 2008).
Increase in soil carbon under trees improves soil physical
properties through forming soil aggregates, or as source of food for
soil fauna thus increasing their activities in the soil. These
processes lead to lower soil density and higher soil porosity as
well as higher water infiltration in the presence of trees as opposed
to bare soils (Bargués et al., 2014; Hansson, 2006; Sanou et al.,
2010; Zomboudré, 2009). As organic matter is the source of
nutrients through its decomposition, higher chemical fertility is
also common in the presence of trees. The trees, by improving the
micro-climate in their influence zone, and increasing the
accumulation of soil organic matter create more favorable
conditions for the micro-organisms that thrive in the rooting
zone of trees in comparison with the open area (INERA, 2000). For
31
example, Diedhiou et al. (2009) reported that Guiera senegalensis and
Piliostigma reticulatum shrubs were found to promote fungi,
microbial diversity and litter decomposition. The high presence
of microorganisms in the rooting zone of trees might partly be due to
the fact that phosphorus, a limiting resource in the soils of the Sahel,
was found to be in a form more accessible under trees (F. albida and V.
paradoxa) compared to the open field (Gnankambary et al., 2008).
In addition to these improved soil conditions, trees reduce
radiation underneath leading to less evapo-transpiration from the
soil and the associated crops under the trees. Another process
leading to increased soil water content under trees is the hydraulic
lift or water redistribution, which is the passive movement of
water through the roots of trees, from deeper and wetter soil layers
to shallower and drier horizons, along a gradient of soil water
potential (Richards and Caldwell, 1987). These have been
investigated and proven in Azadirachta indica in Niger (Smith
et al., 1997), in F. albida in Burkina Faso (Roupsard et al., 1999), in V.
paradoxa and P. biglobosa in Burkina Faso (Bayala et al., 2008a), and
in G. senegalensis and P. reticulatum in Senegal (Kizito et al., 2012).
This process redistributes water to shallow soil layers where it can
be taken up by the plants, enhance transpiration and ultimately
increase biomass production.
One question, “why then associated crops are not always
performing better in such a high potential niche provided by trees
in mixed agricultural systems?” One explanation given is that this
may be related to the layers of soil exploited by the two
components (Bayala et al., 2004; Jones et al., 1998; Lehmann
et al., 1998; Schroth, 1995; Tomlinson et al., 1998). In this situation
both the tree and the crop are sharing the same pool for nutrients
and water, and it has been found that the competition for these
resources causes the poor performance of the less competitive
component, which in most cases are the associated crops (Bayala
et al., 2008c; Kho et al., 2001). However, in complementary
combinations of trees and crops, niche differentiation takes place
whereby the roots of the tree component exploit the lower soil
horizon, leaving the upper soil horizon to associated crops; this
could be related to the type of crop associated with the trees
(Sanou et al., 2012a,b,b).
Light saturation point (point at which increase in light does not
result in increase in photosynthesis) is higher for C4 than C3 crops
(Fig. 1) and that explains their differential responses to light
reduction under tree shade (Osborne et al., 2008). For example,
Sanou et al. (2012b) found higher radiation and water use
efficiencies for P. glaucum (millet), a C4 crop (20.3  2.45 mmol
CO2 mol 1 PAR and 4.8  0.45 mmol CO2 mol 1 H2O, respectively),
compared to the values obtained in C. esculenta (taro), a C3 crop
(14.7  1.15 mmol CO2 mol 1 PAR and 2.3  0.17 mmol CO2 mol 1
H2O, respectively). This is a consequence of three-fold higher
maximum net photosynthesis value registered in P. glaucum
(91.86 mmol m 2 s 2) compared to that in C. esculenta (26.69 mmol
m 2 s 2). Therefore, poor cereal performance under trees has been
reported by many authors to be due to the reduction in light
transmitted under the trees (Bayala et al., 2002; Kater et al., 1992,b;
Sanou et al., 2012a,b). Other reasons given to explain such
reduction in crop performance include increased humidity in
both soil and ambient air leading to lower plant survival because of
fungal attacks, and inter-specific competition for nutrients at
maturity stage (Kater et al., 1992). On the contrary, the increase in
photorespiration due to the high temperature and water stress,
and light saturation could be the reason for the low dry matter
accumulation of C. esculenta in the open field recorded by Sanou
et al. (2012a), as previously reported in some C3 plants (Hay and
Porter, 2006; Osborne et al., 2008). However, these effects may
vary from one species to another as well as between individuals of
the same species, according to the rainy season (Bazié et al., 2012).
P. biglobosa with a round crown type, projecting more heavy shade,
32 J. Bayala et al. / Agriculture, Ecosystems and Environment 205 (2015) 25–35
was found to be more severe in reducing crop yield as opposed to V.
paradoxa with an upright type of branching that allows light in the
morning and afternoon to reach under the tree. In addition, V.
paradoxa tended to show neutral, to positive, impact on cereal
crops during a poor rainy season (Bayala, 2002; Bayala et al., 2002).
The lack of significant difference in crop performance between the
crops under certain tree species and in the open may be due to the
fact that the negative effect of tree shade must have been
compensated for by improvements in crop temperature and soil
fertility (Bayala et al., 2012; Jonsson et al., 1999). It could also be
due to poor studies, low power for various reasons, etc.
The high rates of tree transpiration may also contribute to the
reduction in crop performance if it were inferred that high rates of
tree transpiration would deplete soil moisture from an area
occupied by the tree roots (Bayala et al., 2004). Indeed, when
pruning is applied to the tree crown, light availability is improved
(Bayala et al., 2002; Bazié et al., 2012) while fine root density is
reduced (Bayala et al., 2004), to match the corresponding reduced
demand from the aboveground part, leading to less water
transpired by pruned individuals (Bayala et al., 2002). As a result,
crop yield increase by 4 to 8-fold was observed under pruned trees
by the same authors. However, such an effect must be transient.
Before such benefits can be realized again there must be a period of
canopy and root expansion that will increase competition with
crops. The dynamics in time matter and this has been well studied.
An alternative argument for light limitation is that, when
cereals are replaced by C3 plants, the yield is less reduced with
only, for instance, 2% for cotton under V. paradoxa (Kater et al.,
1992). The performance of C3 plants can even be improved through
increase in leaf area (Sanou et al., 2012b), and probably in stomatal
density (Onwueme and Johnston, 2000), in the shade, allowing
them to intercept more light. The increase in radiation use
efficiency of C. esculenta reported by Sanou et al. (2012b) may have
helped it to avoid a drastic reduction of net assimilation in shade
conditions, while the opposite happens for cereals (Sanou et al.,
2012b; Zomboudré et al., 2005).
The above detailed description of the interactions reveals their
complexity and modeling has been thought to be a useful tool to
help reaching a better understanding of these interactions faster
and cheaper than a purely empirical approach could do (Bayala
et al., 2008c; Coulibaly et al., 2014; Kho, 2000; van Noordwijk and
Lusiana, 1999). Indeed, modeling can help reducing time, labor, and
funds for establishing experiments to test a hypothesis about
explanatory factors. Therefore, this tool has been used to further
explore the experimental data in soil–tree–crop interactions in
parklands. Simulations were used to identify and isolate factors
that determine productivity in agroforestry systems. For example,
the WaNuLCAS model (van Noordwijk and Lusiana, 1999; van
Noordwijk et al., 2011) was run by Bayala et al. (2008c) and the
results indicated that after light, phosphorus and water were the
most strongly limiting factors for crop production under P.
biglobosa and V. paradoxa. However, results of such simulations
need to be verified experimentally (Bayala et al., 2008c).
Combining light, water and fertilizer factors in an experiment,
Bazie et al. (2012) observed non-significant (23–29%) gains in grain
and dry matter yields due to water supply, variable increases in
grain and straw (8–58%) yields associated with the fertilizer
application and a very strong increase (348–520%) associated with
crown pruning or light increase. In a similar attempt to separate
the effects of various factors, Kho et al. (2001) found 36% higher
millet production under F. albida compared to the open field. That
was due to 200% higher N availability under trees causing 26%
production increase whereas P availability was 30% higher causing
a production increase of 13%. In this Faidherbia system, a negligible
3% decrease in crop production was recorded due to competition
for light and water. Using a spline procedure relative to distance
from tree trunk to plot centers, Boffa et al. (2000) showed that the
positive effect of V. paradoxa on sorghum yield extended over
several canopy radii and density. They recommended that, for
medium-size V. paradoxa trees, tree density could range between
30 and 43 individuals ha 1. The above-mentioned findings that led
to this recommendation may have resulted from an example of
possible design mistake. In effect, the assumption in that study was
that the flat part of the response at distance was the same as the
response with no tree. Using WIMISA (WIndbreak-MIllet SAhel)
model and field data collected in Niger, Mayus (1998) showed that
yields were reduced in the vicinity of the windbreak of Bauhinia
rufescens due to competition for water use at the beginning of the
rainy season. The same author observed that under moderate
water stress, light reduction was of similar or major importance
while a slight increase in yields was due to protection against wind
erosion, improved water availability and wind speed reduction.
Coulibaly et al. (2014) have run WaNuLCAS model to simulate the
effects of different management options (tree densities, tree leaf
pruning, mulching and changes in tree root patterns affecting
hydraulic redistribution) under current and future climates on
sorghum production in parklands. These authors found that under
the current climate conditions, increasing the tree density does not
negatively affect sorghum growth if crown pruning is applied. The
effects of using long cycle sorghum, applying tree pruning and
using tree performing hydraulic function with deep root system
were also simulated in relation with the occurrence of extreme
events like flooding and also drought (Coulibaly et al., 2014). This
exercise revealed that long cycle sorghum could be cultivated on a
long-term basis in agroforestry parklands without providing any
organic or inorganic fertilizers and the sorghum performance will
remain stable as opposed to the short cycle ones. Applying pruning
will also be useful when flooding occurs as a consequence of
changing climate as this operation will reduce shade in situation
where tree growth is accelerated by the availability of water. Tree
species with dynamic canopy and performing hydraulic function
with deep root system were found to be suitable for both flooding
and drought situations but for different reasons. Deeps roots do not
have access to the richer soil horizons and therefore tree canopy
will grow slower even in the presence of excess water (flooding)
and the tree canopy will produce less shade. In drought situation,
deep roots are appropriate to perform water redistribution that
will benefit the associated crops in making water and nutrients
available to them (Coulibaly et al., 2014). Modeling has also some
limitations in understanding the processes. Indeed, models are a
way of understanding the implications of processes we know
sufficiently well to structure and parameterize the models. In turn,
models are able to help in understanding additional processes,
except to the extent that they reveal what we do not know when
simulated and observed data disagree.
Understanding the functioning of these systems is important
but this must lead to practical recommendations for farmers and
that has been the backbone of some of the studies that were geared
to test management options such as pruning (Bayala et al., 2002,
2008c; Bazié et al., 2012; Kessler, 1992) and the use of shade
tolerant crops (Pouliot et al., 2012; Sanou, 2010,b; Sanou et al.,
2012a,b; Sidibé, 2010). Pruning of C. pinnata improved groundnut
biomass production with 2.2 t ha 1 under pruned trees compared
to 1.8 t ha 1 under unpruned trees (Samba, 1997). Pruning of S.
setigera induced better production of millet, sorghum and
groundnut even though the shade of this species had more
depressive impacts on cereals compared to groundnut, which is C3
plant (Bakhoum et al., 2001). Root trenching of P. biglobosa and V.
paradoxa led to yield increase in grain (315 kg ha 1 versus
217 kg ha 1) and straw dry biomass (1639 kg ha 1 versus 1307 kg
ha 1) of S. bicolor (Bayala et al., 2013). The results of this study
indicated that root competition is much more important under V.
 J. Bayala et al. / Agriculture, Ecosystems and Environment 205 (2015) 25–35
paradoxa, compared to the competition for light while for P.
biglobosa both above and belowground competitions seem to be
important (Bayala et al., 2013). Most of the reported pruning
experiments have been of short duration and their effects are
transient to crown and root recovery with time (Bayala et al.,
2008b, 2013). Such experiments have also failed to look at
associated processes (fast decomposition of accumulated organic
matter, changes in soil organism communities, etc.) due to the
removal of the tree crowns.
Although, in the short term, crop yields may be reduced due to
competition, in the long-term, favorable soil conditions (soil
fertility) are provided by parkland trees (Bayala et al., 2014). In
addition, tree species like F. albida, A. digitata, Borassus spp., and C.
pinnata can have positive effects on crop yields as they compete
less with crops (Depommier et al., 1992; Kho et al., 2001; Louppe
et al., 1996; Roupsard et al., 1996, 1999; Samba, 1997; Sanou et al.,
2012a; Yaméogo, 2008).
6. Conclusion and perspectives
To better understand the functioning of parkland systems, a
tremendous effort has been made in recent years to develop
methods and conduct studies to collect field data; this effort has
generated a wealth of knowledge about these systems. Modeling
efforts are still at their infancy but they have generated
information which has helped advance our understanding and
testing of some management options. Despite the major break-
throughs, an over-all understanding of the functioning of these
complex systems is still lacking because no published studies have
addressed all possible interactions that take place simultaneously
in such systems. Hence, research is needed to increase our
understanding of the interactions and tradeoffs, and the way these
are modified by specific socio-ecological settings. Therefore, some
unexplored or insufficiently explored areas still remain, indicating
the need to:
 Continue the efforts of designing sound experiments to address
the methodological difficulties of assessing the park effect;
 Try combining yield mapping with modeling to improve the data
analysis approach;
 Accelerate modeling efforts at different scales (e.g., tree–crop,
field and landscape) to improve our understanding and save time
and resources;
 Pursue long-term research on processes to be able to tailor
interventions to particular contexts;
 Continue investigating the tradeoffs and synergies between and
among the goods and services that trees in agro-ecosystems can
provide;
 Analyze whether parklands are better adapted to climate change
than monocultures and whether they can mitigate it;
 Continue the research on how the systems might respond or be
managed differently in relation to climate change forecasts, and;
 Combine bio-physical and socio-economic research.
While waiting for the results of the above suggested new
investigation areas, existing information on the beneficial effects of
trees, for long-term sustainability, needs to be largely shared with
end-users so that they can widely apply the recommendations
derived so far. This is likely to be an important information for
improving the livelihoods of people living in less-productive agro-
ecological conditions of the Sahel.
Acknowledgements
This part of ICRAF’s research is financed by the CGIAR Research
Programs on Forests, Trees and Agroforestry (FTA). We thank
33
anonymous reviewers of earlier versions of the draft. Finally, we
are grateful to an anonymous person who helped redrawing Fig. 1.
References
Arbonnier, M., 2000. Arbres, Arbustes Et Lianes Des Zones Sèches D'afrique De
L'ouest. CIRAD – Muséum d'Histoire Naturelle, Paris, France.
Augusseau, X., Nikiéma, P., Torquebiau, E., 2006. Tree biodiversity, land dynamics
and farmers’ strategies on the agricultural frontier of south-western Burkina
Faso. Biodivers. Conserv. 15, 613–630.
Bakhoum, C., Samba, N.A.S., Ndour, B., 2001. Sterculia setigera del.: effet sur les
cultures. Ann. Forest Sci. 58, 207–215.
Bargués, T.A., Reese, H., Almaw, A., Bayala, J., Malmer, A., Laudon, H., Ilstedt, U., 2014.
The effect of trees on preferential flow and soil infiltrability in an agroforestry
parkland in semiarid Burkina Faso. Water Resour. Res. 50, 3342–3354.
Bayala, J., 2002. Tree Crown Pruning as a Management Tool to Enhance the
Productivity of Parklands in West Africa. PhD Thesis. University of Wales,
Bangor, UK.
Bayala, J., Balesdent, J., Marol, C., Zapata, F., Teklehaimanot, Z., Ouedraogo, S.J., 2006.
Relative contribution of trees and crops to soil carbon content in a parkland
system in Burkina Faso using variations in natural 13C abundance. Nutr. Cycl.
Agroecosyst. 76, 193–201.
Bayala, J., Bazié, H.R., Sanou, J., 2013. Competition and facilitation-related factors
impacts on crop performance in an agro-forestry parkland system in Burkina
Faso. Afr. J. Agric. Res. 8 (43), 5303–5310.
Bayala, J., Heng, L.K., van Noordwijk, M., Ouedraogo, S.J., 2008a. Hydraulic lift study
in two native tree species of agroforestry parklands of West African dry savanna.
Acta Oecol. 34, 370–378.
Bayala, J., Kindt, R., Belem, M., Kalinganire, A., 2011. Factors affecting the dynamics of
tree diversity in agroforestry parklands for cereal and cotton farming system in
Burkina Faso. New Forest 41, 280–296.
Bayala, J., Lamien, N., Ouédraogo, S.J., 2000. Etat et tendances évolutives du parc à
karité dans le système de production cotonnière de Yasso (Sud-Ouest du
Burkina Faso). Sci. Tech. 24 (2), 89–104.
Bayala, J., Ouedraogo, S.J., Teklehaimanot, Z., 2008b. Rejuvenating indigenous trees
in agroforestry parkland systems for better fruit production using crown
pruning. Agroforest. Syst. 72, 187–194.
Bayala, J., Sanou, J., Teklehaimanot, Z., Kalinganire, A., Ouédraogo, S.J., 2014.
Parklands for buffering climate risk and sustaining agricultural production in
the Sahel of West Africa. Curr. Opin. Environ. Sustain. 6, 28–34.
Bayala, J., Sileshi, G.W., Coe, R., Kalinganire, A., Tchoundjeu, Z., Sinclair, F., Garrity, D.,
2012. Cereal yield response to conservation agriculture practices in drylands of
West Africa: a quantitative synthesis. J. Arid Environ. 78, 13–25.
Bayala, J., Teklehaimanot, Z., Ouedraogo, S.J., 2002. Millet production under pruned
tree crowns in a parkland system in Burkina Faso. Agroforest. Syst. 54, 203–214.
Bayala, J., Teklehaimanot, Z., Ouedraogo, S.J., 2004. Fine root distribution of pruned
trees and associated crops in a parkland system in Burkina Faso. Agroforest.
Syst. 60, 13–26.
Bayala, J., van Noordwijk, M., Lusiana, B., Kasanah, N., Teklehaimanot, Z., Ouedraogo,
S.J., 2008c. Separating the tree–soil–crop interactions in agroforestry parkland
systems in Saponé (Burkina Faso) using WaNuLCAS. Adv. Agroforest. 4, 296–
308.
Bazié, H.R., 2013. Interactions arbres-cultures et rôles des parcs agroforestiers à
Vitellaria paradoxa (C.F. Gertn.) sur les paramètres du bilan hydrique au Burkina
Faso. Thèse de Doctorat Unique. Université de Ouagadougou, Burkina Faso.
Bazié, H.R., Bayala, J., Zombré, G., Sanou, J., Ilstedt, U., 2012. Separating competition-
related factors limiting crop performance in an agroforestry parkland system in
Burkina Faso. Agroforest. Syst. 84, 377–388.
Boffa, J.M., 1995. Productivity and Management of Agroforestry Parklands in the
Sudan Zone of Burkina Faso, West Africa. PhD. Dissertation. Perdue University,
West Lafayette, Indiana, USA.
Boffa, J.M., 1999. Agroforestry Parklands in Sub-Saharan Africa. FAO Conservation
Guide 34, Rome, Italy.
Boffa, J.M., Taonda, S.J.B., Dickey, J.B., Knudson, D.M., 2000. Field-scale influence of
karité (Vitellaria paradoxa) on sorghum production in the Sudan zone of Burkina
Faso. Agroforest. Syst. 49, 153–175.
Bonkoungou, E.G., Ayuk, E.T., Depommier, D., Zoungrana, I., 1997. Les parcs
agroforestiers des zones semi-arides d'Afrique de l'Ouest. Actes Du Séminaire
International. ICRAF/IRBET/CILSS/LTC, Ouagadougou, Burkina Faso 25–27
Octobre 1993, ICRAF, Nairobi.
Breman, H., Kessler, J.J., 1995. Woody Plants in Agro-Ecosystems of Semi-Arid
Regions (With an Emphasis on the Sahelian Countries). Springer-Verlag, Berlin.
Brouwer, J., Fussell, L.K., Herrmann, L., 1993. Soil and crop growth micro-variability
in the West African semi-arid tropics: a possible risk-reducing factor for
subsistence farmers. Agric. Ecosys. Environ. 45, 229–238.
Bruno, J.F., Stachowicz, J.J., Bertness, M.D., 2003. Inclusion of facilitation into
ecological theory. Trends Ecol. Evol. 18, 119–125.
Charreau, C., Vidal, P., 1965. Influence de l'Acacia albida del. sur le sol: la nutrition
minérale et les rendements des mils Pennisetum au Sénégal. Agron. Trop. 6–7,
660–686.
Coulibaly, Y.N., Mulia, R., Sanou, J., Zombré, G., Bayala, J., Kalinganire, A., van
Noordwijk, M., 2014. Crop production under different rainfall and management
conditions in agroforestry parkland systems in Burkina Faso: observations and
simulation with WaNuLCAS model. Agroforest. Syst. 88, 13–28.
34 J. Bayala et al. / Agriculture, Ecosystems and Environment 205 (2015) 25–35
Dembélé, D., 1994. Ecophysiologie de Faidherbia albida, sa répartition et son effet
agronomique. Mémoire Ingénieur IDR, Option Eaux et Forêts. Université de
Ouagadougou, Burkina Faso.
Depommier, D., Janodet, E., Roupsard, O., 1992. Faidherbia albida parks and their
influence on soils and crops at Watinoma, Burkina Fas. In: Vandenbeldt, R.J.
(Ed.), Faidherbia Albida in the West African Semi-arid Tropics. Proceedings of a
Workshop, 22–26 April 991. ICRISAT, Niamey, Niger, pp. 111–118.
Diedhiou, S., Dossa, E.L., Badiane, A.N., Diedhiou, I., Sene, M., Dick, R.P., 2009.
Decomposition and spatial microbial heterogeneity associated with native
shrubs in soils of agroecosystems in semi-arid Senegal. Pedobiologia 52, 273–
286.
Dossa, E.L., Diedhiou, I., Khouma, M., Sene, M., Lufafa, A., Kizito, F., Samba, S.A.N.,
Badiane, A.N., Diedhiou, S., Dick, R.P., 2012. Crop productivity and nutrient
dynamics in a shrub (Guiera senegalensis)-based farming system of the Sahel.
Agron. J. 105, 1255–1264.
Ellis, T., 2013. Agroforestry-livestock systems research in West Africa. An overview
of work undertaken 2011-2013 in the AFSI project CerLiveTrees (CLT) funded by
CORAF-CSIRO and foreshadowed for future tree system research in Phase 3 of
AFSI - July 2013.
Faye, M.D., Weber, J.C., Abasse, T.A., Boureima, M., Larwanou, M., Bationo, A.B.,
Diallo, B.O., Sigué, H., Dakouo, J.-M., Samaké, O., Sonogo-Diaité, D., 2011.
Farmers’ preferences for tree functions and species in the West African Sahel.
Forest Trees Livelihoods 20, 113–136.
Food and Ariculture Organization (FAO), 1992. Legume trees and other fodder trees
as protein sources for livestock. In: Speedy, A., Pugliese, P.L. (Eds.), Proceedings
of the FAO Expert Consultation held at the Malaysian Agricultural Research and
Development Institute (MARDI) in Kuala Lumpur, Malaysia, 14–18 October 1991.
FAO, Rome, Italy.
Furbank, R.T., von Caemmerer, S., Sheehy, J., Edwards, G., 2009. C4 rice: a challenge
for plant phenomics. Funct. Plant Biol. 36, 845–856.
Geiger, S.C., Brown, K.W., Vandenbeldt, R.J., 1994. Variability in the growth of
Faidherbia albida: the soils connection. Soil Sci. Soc. Am. J. 58, 227–231.
Ghannoum, O., 2009. C4 photosynthesis and water stress. Ann. Bot. 103 (4), 635–
644.
Gijsbers, H.J.M., Kessler, J.J., Knevel, M.K., 1994. Dynamics and natural regeneration
of woody species in farmed parkland in the Sahel region (Province of Passoré)
Burkina Faso. Forest Ecol. Manag. 64, 1–12.
Gnankambary, Z., Ilstedt, U., Nyberg, G., Hien, V., Malmer, A., 2008. Nitrogen and
phosphorus limitation of soil microbial respiration in two tropical agroforestry
parklands in the south-Sudanese zone of Burkina Faso: the effects of tree
canopy and fertilization. Soil Biol. Biochem. 40, 350–359.
Gonzalez, P., 2001. Desertification and a shift of forest species in the West African
Sahel. Clim. Res. 17, 217–228.
Gonzalez, P., Tucker, C.J., Sy, H., 2012. Tree density and species decline in the African
Sahel attribute to climate. J. Arid Environ. 78, 55–64.
Gowik, U., Westhoff, P., 2011. The path from C3 to C4 photosynthesis. Plant Physiol.
155, 56–63.
Hadgu, K.M., Rossing, W.A.H., Kooistra, L., van Bruggen, A.H.C., 2009. Spatial
variation in biodiversity, soil degradation and productivity in agricultural
landscapes in the highlands of Tigray, northern Ethiopia. Food Sec. 1, 83–97.
Hansen, H.H., Sanou, L., Nacoulma, B.M.I., 2008. Tree leaves in the diet of free-
ranging ruminants in three areas of Burkina Faso. Livest. Res. Rural Dev. 20 (3),
33. http://www.lrrd.org/lrrd20/3/hann20033.htm.
Hansson, L., 2006. Comparisons of infiltration capacities in different parklands and
farming systems of semi-arid Burkina Faso. MSc Thesis. Swedish University of
Agricultural Sciences, Department of Forest Ecology, Sweden.
Hay, R., Porter, J., 2006. The Physiology of Crop Yield. Blackwell, Oxford (UK).
INERA, 2000. Minimizing competition in dry land agroforestry. INCO-DC:
International Cooperation with Developing Countries (1994-1998). Second
Annual Report for the Period of 1-11-1999 to 31-10-2000. INERA, Ouagadougou,
Burkina Faso.
Johnston, M., Onwueme, I.C., 1998. Effect of shade on photosynthetic pigments in
the tropical root crops: Yam, Taro, Tannia, Cassava and Sweet Potato. Exp. Agric.
34, 301–312.
Jones, M., Sinclair, F.L., Grime, V.L., 1998. Effects of tree species and crown pruning on
root length and soil water content in semi-arid agroforestry. Plant Soil 201, 197–
207.
Jonsson, K., 1995. Agroforestry in dry savanna areas in Africa: interactions between
trees, soils and crops. PhD Dissertation. Swedish University of Agricultural
Sciences, Umea, Sweden.
Jonsson, K., Ong, C.K., Odongos, J.C.W., 1999. Influence of scattered néré and karité
on microclimate: soil fertility and millet yield in Burkina Faso. Exp. Agric. 35,
39–53.
Kater, L.J., Kante, S., Budelman, A., 1992. Karité (Vitellaria paradoxa) and néré (Parkia
biglobosa) associated with crops in South Mali. Agroforest. Syst. 18, 89–105.
Kelty, M.J., Cameron, I.R., 1995. Plot designs for analysis of species interactions in
mixed stands. Commonw. Forest Rev. 74, 322–332.
Kessler, J.J., 1992. The influence of karité (Vitellaria paradoxa) and néré (Parkia
biglobosa) trees on sorghum production in Burkina Faso. Agroforest. Syst. 17, 97–
118.
Kho, R.M., 2000. A general tree–environment–crop interaction equation for
predictive understanding of agroforestry systems. Agric. Ecosys. Environ. 80,
87–100.
Kho, R.M., Yacouba, B., Yayé, M., Katkoré, B., Moussa, A., Iktam, A., Mayaki, A., 2001.
Separating the effects of trees on crops: the case of Faidherbia albida and millet
in Niger. Agroforest. Syst. 52 (3), 219–238.
Kizito, F., Dragila, M.I., Sene, M., Brooks, J.R., Meinzer, F.C., Diedhiou, I., Diouf, M.,
Lufafa, A., Dick, R.P., Selker, J., Cuena, R., 2012. Hydraulic redistribution by two
semi-arid shrub species: implications for Sahelian agro-ecosystems. J. Arid
Environ. 83, 69–77.
Lamien, N., Sidibé, A., Bayala, J., 1996. Use and commercialization of non-timber
forest products in western Burkina Faso. In: Leakey, R.R.B., Temu, A.B., Melnyk,
M. (Eds.), Domestication and Commercialization of Non-timber Forest Products
in Agroforestry Systems. FAO, Rome Italy, pp. 51–63 Non-wood Forest Products
9.
Lehmann, J., Peter, I., Steglich, C., Gebauer, G., Huwe, B., Zech, W., 1998. Below-
ground interactions in dryland agroforestry. Forest Ecol. Manag. 111, 157–169.
Le Houerou, H.N., 1980. Browse in Africa: The Current State of Knowledge. I.L.C.A,
Addis Ababa, Ethiopia.
Louppe, D., N'Dour, B., Samba, A.N.D., 1996. Influence de Faidherbia albida sur
I'arachide et le mil au Sénégal. Méthodologie de mesure et estimations des
effets d'arbres émondés avec ou sans parcage d'animaux, in: Peltier, R. (Ed.) Les
parcs à Faidherbia, CIRAD-Forêt, Cahiers Scientifiques N 12, Montpellier, pp.
123–139.
Maïga, A., 1987. L'arbre dans les systèmes agroforestiers traditionnels dans la
province du Bazèga. Influence du karité, du néré et de Acacia albida sur le sorgho
et le mil. Rapport de stage. IRBET/CNRST, Ouagadougou, Burkina Faso.
Maranz, S., 2009. Tree mortality in the African Sahel indicates an anthropogenic
ecosystem displaced by climate change. J. Biogeogr. 36, 1181–1193.
Mayus, M., 1998. Millet growth in windbreak-shielded fields in the Sahel. PhD
Thesis. Wageningen University, The Netherlands.
Mochida, K., Shinozaki, K., 2010. Genomics and Bioinformatics resources for crop
improvement. Plant Cell Physiol. 51 (4), 497–523.
Moussa, H., 1997. Germination du palmier doum (Hyphaene thebaica Mart.) et
analyse de son interaction avec le mil (Pennisetum glaucum L.) en zone semi-
aride du Niger. PhD Université de Laval, Québec, Canada.
Ndiaye, M., Ganry, F., Oliver, R., 2000. Alley cropping of maize and Gliricidia sepium in
the Sudanese Sahel region: some technical feasibility aspects. Arid Soil Res.
Rehab. 14, 317–327.
Nur Osman, A., Ræbild, A., Christiansen, J.L., Bayala, J., 2011. Performance of cowpea
(Vigna unguiculata) and pearl millet (Pennisetum glaucum) intercropped under
Parkia biglobosa in an agroforestry system in Burkina Faso. Afr. J. Res. 6 (4), 882–
891.
Ong, C.K., Leakey, R.R.B., 1999. Why tree–crop interactions in agroforestry appear at
odds with tree–grass interactions in tropical savannahs. Agroforest. Syst. 45,
109–129.
Onwueme, I.C., Johnston, M., 2000. Influence of shade on stomatal density, leaf size
and other characteristics in the major tropical root crops, tannia, sweet potato,
yam, cassava and taro. Exp. Agric. 36, 509–516.
Osborne, C.P., Wythe, E.J., Ibrahim, D., Gilbert, M.E., Ripley, B.S., 2008. Low
temperature effects on leaf physiology and survivorship in the C3 and C4
subspecies of Alloteropsis semialata. J. Exp. Bot. 59 (7), 1743–1754.
Ouédraogo, S.J., 1995. Les parcs agroforestiers au Burkina Faso. Rapport de
consultation IRBET/CNRST Ouagadougou. No 79.
Pouliot, M., Bayala, J., Ræbild, A., 2012. Testing the shade tolerance of selected crops
under Parkia biglobosa (Jacq.) Benth. in an agroforestry parkland in Burkina Faso,
West Africa. Agroforest. Syst. 85, 477–488.
Prudencio, Y.C., 1983. Ring management if soils and crops in West African semi-arid
tropics: the case of the mossi farming system in Burkina Faso. Agric. Ecosyst.
Environ. 47, 237–264.
Pullan, R.A., 1974. Farmed parkland in West Africa. Savanna 3 (2), 119–151.
Ræbild, A., Hansen, U.B., Kambou, S., 2012. Regeneration of Vitellaria paradoxa and
Parkia biglobosa in a parkland in southern Burkina Faso. Agroforest. Syst. 85,
443–453.
Raison, J.P., 1988. Les parcs en Afrique: état des connaissances, perspectives de
recherches. Document de travail, Centres d'Etudes Africaines, EHESS, Paris.
Richards, J.H., Caldwell, M.M., 1987. Hydraulic lift: substantial nocturnal water
transport between layers by Artemisia tridentata roots. Oecologia 73,
486–489.
Ripley, B.S., Abraham, T.I., Osborne, C.P., 2008. Consequences of C4 photosynthesis
for the partitioning of growth: a test using C3 and C4 subspecies of Alloteropsis
semialata under nitrogen-limitation. J. Exp. Bot. 59 (7), 1705–1714.
Ripley, B.S., Gilbert, M.E., Ibrahim, D.G., Osborne, C.P., 2007. Drought constraints on
C4 photosynthesis: stomatal and metabolic limitations in C3 and C4 subspecies
of Alloteropsis semialata. J. Exp. Bot. 58 (6), 1351–1363.
Roupsard, O., Depommier, D., Janodet, E., 1996. Influence de Faidherbia albida sur le
sol et le sorgho: Observations dans le parc de Watimona au Burkina Faso, in:
Peltier, R. (Ed.) Les parcs à Faidherbia, CIRAD-Forêt, Cahiers Scientifiques N 12,
Montpellier, pp. 141–152.
Roupsard, O., Ferh, A., Granier, A., Pallo, F., Depommier, D., Mallet, B., Joly, H.J.,
Dreyer, E., 1999. Reverse phenology and dry-season water uptake by Faidherbia
albida (Del.) A. Chev. in an agroforestry parkland of Sudanese West Africa. Funct.
Ecol. 13, 450–472.
Sabiitti, E.N., Cobbina, J., 1992. Parkia biglobosa: a potential multipurpose fodder
tree legume in West Africa. Int. Tree Crops J. 7, 113–139.
Samba, S.A.N., 1997. Influence de Cordyla pinnata sur la fertilité d'un sol ferrugineux
tropical et sur le mil et l'arachide dans un système agroforestier traditionnel au
Sénégal. Thèse de PhD Faculté de Foresterie et de Géomatique, Université Laval,
Québec, Canada.
Sanon, H.O., Kaboré-Zoungrana, C., Ledin, I., 2007. Behaviour of goats, sheep and
cattle and their selection of browse species on natural pasture in a Sahelian area.
Small Rumin. Res. 67, 64–74.
 J. Bayala et al. / Agriculture, Ecosystems and Environment 205 (2015) 25–35
Sanou, J., 2010. Optimizing Productivity of Agroforestry Parkland Systems in West
Africa Using Shade Tolerant Annual Crops. PhD Thesis. University of Wales,
Bangor, UK.
Sanou, J., Bayala, J., Bazié, P., Teklehaimanot, Z., 2012a. Photosynthesis and biomass
production by millet (Pennisetum glaucum) and taro (Colocasia esculenta) grown
under baobab (Adansonia digitata) and néré (Parkia biglobosa) in an agroforestry
parkland system of Burkina Faso (West Africa). Exp. Agric. 48 (2),
283–300.
Sanou, J., Bayala, J., Teklehaimanot, Z., Bazie, P., 2012b. Effect of shading by baobab
(Adansonia digitata) and néré (Parkia biglobosa) on yields of millet (Pennisetum
glaucum) and taro (Colocasia esculenta) in parkland systems in Burkina Faso,
West Africa. Agroforest. Syst. 85, 431–441.
Sanou, J., Zougmoré, R., Bayala, J., Teklehaimanot, Z., 2010. Soil infiltrability and
water content as affected by Baobab (Adansonia digitata L.) and Néré (Parkia
biglobosa (Jacq.) Benth.) trees in farmed parklands of West Africa. Soil Use
Manag. 26, 75–81.
Schroth, G., 1995. Tree root characteristics as criteria for species selection and
systems design in agroforestry. Agroforest. Syst. 30, 125–143.
Sidibé, D., 2010. Improving the Management and Productivity of Ziziphus Mauritiana
and Tamarindus Indica in Agroforestry Parkland Systems in Mali, West Africa.
PhD Thesis. School of Environment, Natural Resources and Geography, Bangor
University, UK.
Smith, D.M., Jarvis, P.G., Odongo, J.C.W., 1997. Sources of water used by trees and
millet in Sahelian windbreak systems. J. Hydrol. 198, 140–153.
Takimoto, A., Nair, P.K.R., Nair, V.D., 2008. Carbon stock and sequestration potential
of traditional and improved agroforestry systems in the West African Sahel.
Agric. Ecosys. Environ. 125, 159–166.
Tilander, Y., Ouedraogo, G., Yougma, F., 1995. Impact of tree coppicing on tree–crop
competition in parkland and alley farming systems in semiarid Burkina Faso.
Agroforest. Syst. 30, 363–378.
Tomlinson, H., Traoré, A., Teklehaimanot, Z., 1998. An investigation of root
distribution of Parkia biglobosa in Burkina Faso, West Africa, using a logarithmic
spiral trench. Forest Ecol. Manag. 107, 173–182.
35
van Noordwijk, M., Lusiana, B., 1999. WaNuLCAS a model of water, nutrient and light
capture in agroforestry systems. Agroforest. Syst. 43, 217–242.
van Noordwijk, M., Lusiana, B., Khasanah, N., Mulia, R., 2011. WaNuLCAS Version 4.0,
Background on a Model of Water, Nutrient and Light Capture in Agroforestry
Systems. International Center for Research in Agroforestry (ICRAF), Bogor,
Indonesia.
van Noordwijk, M., Ong, C.K., 1999. Can the ecosystem mimic hypothesis be applied
to farms in African savannahs? Agroforest. Syst. 45, 131–158.
van Noordwijk, M., Bayala, J., Hairiah, K., Lusiana, B., Muthuri, C., Khasanah, N.,
Mulia, R., 2014. Agroforestry Solutions for Buffering Climate Variability and
Adapting to Change. In: Fuhrer, J., Gregory, P.J. (Eds.), Climate change Impact and
Adaptation in Agricultural Systems, , pp. 216–232.
von Maydell, H.J., 1983. Arbres Et Arbustes Du Sahel. Leurs Caractéristiques Et Leurs
Utilisations. GTZ, Eschborn, Germany.
Wezel, A., Lykke, A.M., 2006. Woody vegetation change in Sahelian Africa: evidence
from local knowledge. Environ. Dev. Sustain. 8, 553–567.
Wilson, T.D., Brook, R.M., Tomlinson, H.F., 1998. Interactions between néré (Parkia
biglobosa) and under-planted sorghum in parkland system in Burkina Faso. Exp.
Agric. 35, 85–98.
Yaméogo, J., 2008. Contribution des parcs à Borassus akeasii Bayton, Ouédraogo et
Guinko au fonctionnement des systèmes de productions dans le sud-ouest du
Burkina Faso. Thèse Unique. Université de Ouagadougou, Burkina Faso.
Zomboudré, G., 2009. Caractérisation biophysique et incidence des parcs à Vitellaria
paradoxa Gaertn. et Faidherbia albida (Del.) A. Chev. Sur les facteurs
pédoclimatiques et la productivité du maïs (Zea mays L.) dans la zone ouest du
Burkina Faso. Thèse Doctorat Unique. Université de Ouagadougou, Burkina Faso.
Zomboudré, G., Zombré, G., Ouedraogo, M., Guinko, S., Macauley, H.R., 2005.
Réponse physiologique et productivité des cultures dans un système
agroforestier traditionnel: cas de maïs (Zea mays L.) associé au karité (Vitellaria
paradoxa Gaertn.) dans la zone est du Burkina Faso. BASE 9, 75–85.
Zomer, R.J., Trabucco, A., Coe, R., Place, F., 2009. Trees on farm: analysis of global
extent and geographical patterns of agroforestry. Working Paper No. 89. The
World Agroforestry Centre, Nairobi, Kenya.