AGRICULTURA TROPICA ET SUBTROPICA VOL.

43 (4) 2010

Review Articles

THE USE AND INTEGRATION OF INGA EDULIS IN AGROFORESTRY SYSTEMS

IN THE AMAZON – REVIEW ARTICLE

LOJKA B.1, DUMAS L.2, PREININGER D.1, POLESNY Z.1, BANOUT J.1

1
Institute of Tropics and Subtropics, Czech University of Life Sciences, Czech Republic
2
Ecole Nationale Supérieure Agronomique de Montpellier, Montpellier, France

Abstract

In Latin America this fast growing, acid-soil tolerant tree, which improves soil fertility through nitrogen Þxation, is tradi-
tionally used to shade perennial crops such as coffee and cacao, provide Þrewood and charcoal, and produce a sweet pulp
suitable for human consumption. I. edulis is also useful as a green manure with its high biomass production and helps control
weeds and erosion, in alley cropping and other agroforestry systems. It has a great potential for restoring degraded soils as
a part of agroforestry system. The information about the physiology, the use by local farmers, the silvicultural management
and integration into agroforestry systems were collected. The species is commonly used in a large variety of agroforestry
systems, however, it is surprising that it is still not the object of any improved breeding program and scientiÞc information
about this species are still rather scarce.

Key words: Peruvian Amazon, slash-and-burn farming, small farmers, weed control

INTRODUCTION species for agroforestry and other environmental improv-

ing systems (Weber et al., 2001).
The genus Inga (Leguminosea: Mimosoideae: Ingeae) Several researchers have described I. edulis as an agro-
comprises c. 300 species of trees restricted to tropical forestry tree with an important role for farmers and with
America. It is and ubiquitous component of lowland and a great adaptability for agroforestry systems (Szott 1987;
montane rainforest throughout the humid tropical zones Alegre 1991; Salazar and Palm 1991; Fernandes et al.,
from Mexico to Uruguay. The highest species diversity 1991; Ricse and Szott, 1993; Flores 1997; Weber et al.,
is concentrated it the Andean foothills of Peru, Ecuador, 1997; Labarta and Weber 1998; Pennington and Fern-
Colombia and in southern Central America (Pennington, andes, 1998). In the study of Labarta and Weber (1998)
1997). Inga edulis Mart. is one of the most widely dis- of economic valuation of agroforestry species in the Pe-
tributed and economically useful in the whole Amazon ruvian Amazon, I. edulis was ranked as a second most
region. Sotelo-Montes and Weber (1997) investigated important trees species used by local farmers. In other
farmers’ preferences for agroforestry tree species in the study of agroforestry species prioritization (Villachica,
Peruvian Amazon and identiÞed I. edulis as a high pri- 1995) I. edulis was generally cited as a tree with facili-
ority species for agroforestry research and development. ties to integrate agroforestry systems and with excellent
This fast growing, acid-soil tolerant tree, which improves characteristics to develop its use in agroforestry. The ob-
soil fertility through nitrogen Þxation, is traditionally jective of this review was to collect most relevant infor-
used to shade perennial crops such as coffee and cacao, mation about the integration and use of this species in
provide Þrewood and charcoal, and produce a sweet pulp agroforestry systems.
suitable for human consumption (Pennington and Fern-
andes, 1998). I. edulis is also useful as a green manure Physiology and adaptation
with its high biomass production and helps control weeds
and erosion (Lawrence, 1995). It has a great potential for The native range of I. edulis is in Amazonian Bra-
restoring degraded soils as a part of agroforestry system. zil, Bolivia, Peru, Ecuador and Colombia. The species
Its presence throughout the Amazon basin sparked inter- has also been introduced across most of tropical South
est of scientists in further exploring the potential of this America, Panama and Costa Rica (Lawrence, 1993). It

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grows in hot, humid climates between 26°S and 10°N, per cm3 it is used mainly as a fuel wood but also for rural
and up to 2000 m elevation and it is most wide-spread in construction.
areas without a dry season or with a dry season of three
to four months and annual rainfall of around 1500 mm The use
(900–3500 mm). It can tolerate short droughts, although
in its natural range some rain falls every month. I. edulis has a wide range of uses among small farmers
I. edulis thrives on all the kind of soils presented in in Latin America (Pennington and Fernandes, 1998). Its
Amazon from fertile alluvial Entisols to strongly-acid, major products are fruit and Þrewood, and its major serv-
infertile Ultisols. Sanchez (1987) described I. edulis as ice functions are shade and soil improvement. Its com-
an excellent species for acid soils in the tropics. How- mon uses are as a mulch supply in alley cropping system,
ever, its growth is better on the non-ßooding Þelds a pasture or soil improver, a shade for perennial crops;
(TCA, 1997). Rapid growth of this species is also re- its potential is as a fuelwood and timber producer and in
ported on highly acidic soil, low in nutrients and high in improved fallows. Other uses are presented in the litera-
aluminum toxicity of abandoned pasture in Amazonian ture such as living fence or medicine properties (Weber
Ecuador (Neill and Nixon, 1998). Leblanc et al. (2005) et al., 1997). The seeds are valued as a natural purgative
have studied the N2-Þxing symbiosis with I. edulis. The for human and cattle.
four strains of symbiotic bacteria were closely related to The pods of Inga edulis are widely marketed and eaten
Bradyrhizobium japonicum and to Bradyrhizobium liao- as a fresh fruit in the Amazon. The fruits of low quality
ningense. The estimate of the percentage of N Þxed from are consumed by the livestock, pigs, poultry or Þshes.
atmosphere out of total N in I. edulis seedlings was about The important content of proteins in the seed confers
40% in the sand media treatments and 10% in the native the potential as a source of animal alimentation (TCA,
soil. 1997). The pulp covering the seeds is sweet and tender,
Tree can reach a height of 30m and a stem diameter contains over 80% water, is rich in carbohydrates and
(dbh) of 60 cm, but usually grows much lower (up to contains a lot of energy. It is eaten usually fresh but juice
15 m) and branches from below 1 m. The branches form can be prepared and it has also a potential for produc-
a broad, ßat and moderately dense canopy. The bark is ing alcoholic beverages. The nutritive value of fresh pulp
pale grey and smooth, with pale elongated lenticels. The is low, but the embryos of this and other Inga species
young twigs are angular in cross-section and covered in can be cooked and are more nutritious than the fruit.
line short brown hair. The leaves are once pinnate, up to The cooking probably degrades trypsin inhibitors and
24 cm long, with four to six pairs of opposite leaßets. enhances palatability (Leakey, 1997). The fruits, which
The terminal pair of leaßets is larger than the basal pair can normally be kept for only three to four days, can be
and can be up to 18 cm long and 11 cm wide. Between stored in a refrigerator for three weeks.
each leaßet there is a nectary gland on the leaf rachis. Concerning the use of I. edulis as a wood source, it is
The inßorescences are dense axillaries spikes of ßowers, generally appreciated for three reasons (1) it is a fast-
each consisting of a calyx tube with 5 lobes (4–9 mm growing tree, (2) its management is easy, and (3) it has
long), a corolla tube with Þve lobes (13–25 mm long) a short fruit production period (6–10 years). The wood
and a large number of white stamens up to 4.5 cm long is used as fuelwood or in the rural construction. (TCA,
united in a tube in the lower half (Pennigton, 1997). 1997). It has a high caloriÞc content and its weak smoke
In observations made in Brazilian Amazon (Falcao and derangement, however, the species is not usually culti-
Clement, 2000) the majority of plants had four ßowering vated speciÞcally for fuelwood or other woody products
periods during the year, with ßowering peaks in March, (Lawrence, 1993).
May, August/September, October/December, with some Leblanc et al. (2005) have described I. edulis as tree
trees presenting Þve ßowering peaks, followed by fruit- used traditionally for shade in coffee and cacao planta-
ing periods in April, June, September/October, and No- tions throughout tropical America. Indeed it is one of the
vember/January. Three to four year old trees produced most popular shade trees for coffee and cacao planta-
20 000–100 000 ßowers (mean 50 000) and 200–800 tions in the humid Neotropics. In Costa Rica, I. edulis is
fruits (mean 500). Fruit set varied from 0.4 to 1.8%, chosen in preference to Erythrina poeppigiana (another
with a mean of 1.1%. Fruit is a large, cylindrical, ribbed commonly used species for coffee shade trees) because it
pod (40–180 cm long), Þlled with 2–3 cm long, usually retains its leaves in the dry season, while E. poeppigiana
black, oval seeds, which are covered with white, cottony, does not (Nichols, 1990). In pastoral systems, I. edulis
edible pulp. Fruit weight varied from 250 to 600 g, with can provide shade for livestock (Weber et al., 1997).
only 22 ± 4% of edible pulp (Falcao and Clement, 2000). Because of the Inga edulis use as a soil fertility im-
The wood has white-grey colour, with density of 0.54 g provement, the production of biomass is an important

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Tab. 1: Potential of biomass growth (in dry matter) of I. edulis

Total above-
Age
Trees Prunning Spacing DM of leaves DM of wood DM of leaves DM of wood ground
of tree
per ha (%) (m) (kg/tree) (kg/tree) (Mg/ha) (Mg/ha) biomass
(years) (Mg/ha)
1 5000 0 1×2 0.4 0.3 2.10 1.65 3.75
2 2500 50 2×2 1.1 2.4 2.80 6.08 8.88
3 1250 75 2×4 3.0 17.7 3.78 22.11 25.89
4 625 88 4×4 4.3 35.9 2.69 22.43 25.12
5 313 94 4×8 5.8 65.4 1.82 20.48 22.29
6 156 97 8×8 6.1 73.0 0.95 11.39 12.34

characteristic. Lojka et al. (2005) have studied the corre- the relatively large recalcitrant mulch fraction, the litter
lation of aboveground biomass growth with other growth may promote carbon sequestration and long-term N ac-
parameters in Peruvian Amazon. Using statistical tools, cumulation in soil.
they have determined that the diameter of the trunk is Because of these N-Þxation characteristics, I. edulis
a more useful indicator of growth and biomass produc- is sometimes used to improve the soil fertility during
tion. Highest correlation for wood biomass was obtained fallows or in alley-cropping systems. Szott and Palm
for dependence between wood biomass and diameter at (1996) have studied the nutrient stocks in I. edulis fal-
10 cm above soil surface: y = 0.0466 x2.3713; R2 = 0.924. lows in Peruvian humid tropics during 53 months on a
Also for leaf biomass high correlation was obtained for previously cultivated Ultisol (Acrisol). They concluded
dependence on the trunk diameter: y = 0.1564 x1.1817; R2 that total N stocks increased by 10% in the I. edulis treat-
= 0.792. Using these equations with high correlation the ment. Total stocks of P and K were 40–80% and 12%
potential aboveground biomass growth for I. edulis can greater than initial values, but Ca and Mg stocks were
be assessed (Table 1.). reduced by 25–40%. Although there were net decreases
The highest biomass production rates reported in the of stocks of P, K, Ca and Mg in soil in all treatments
literature (Szott et al., 1995) reached as much as 31 Mg. during the fallow, storage of P and K in vegetation and
ha–1.yr–1on acid soils at experimental station near Yurim- litter in the I. edulis fallows offset losses of these nu-
aguas, Peru. By two years of age, average tree diameter trients from soil. These results suggest that leguminous
at 10 cm above soil surface ranged up to 10.7 cm and av- biomass of I. edulis may increase N, P and K stocks, but
erage tree height up to 9 m. However, even at lower pro- that incomplete recuperation of Ca and Mg may limit the
duction levels of 8–12 Mg.ha–1.yr–1, I. edulis produced sustainability of short-rotation fallow-based systems on
more biomass than other non-Inga species (Lawrence et acidic, infertile soils.
al., 1995). According to the results of modelling study made by
Oglesby and Fownes (1992) describe the patterns of Lojka et al. (2008) in Peruvian Amazon, improved fal-
I. edulis leaves decomposition. Cumulative N minerali- low with I. edulis has potential in increasing nitrogen
zation was negatively correlated with polyphenol con- levels in the soil, slightly increasing soil organic matter
tent for earlier time periods (1 through 8 weeks) and content, and maintaining soil phosphorus content com-
with lignin for later time periods (4 through 12 weeks). pared to natural bush fallow.
Initial percent N and lignin/N ratio were not strongly Bishop (1983) describes a system developed in Ama-
correlated with N mineralization. The best chemical zonian Ecuador in which the fallow period in shifting
index of N release was the initial polyphenol/N ratio. cultivation sequences was intensiÞed by the use of the
General characteristics about I. edulis green mulch de- Asian perennial forage cover crop Desmodium ovalifo-
composition are also given by Leblanc et al. (2005). lium under I. edulis. In this case, both species improve
Hemicelluloses disappeared almost completely from soil fertility and reduce erosion. Alegre and Rao (1995)
the litter during the 20-week incubation period, while had studied the soil and water conservation by contour
no signiÞcant lignin decay occurred. After a slow start, hedging of I. edulis in the Peruvian humid tropics. They
cellulose partially decayed following linear kinetics. concluded that contour hedgerow intercropping is rec-
The half-life of labile N, estimated as a Michaelis- ommended for moderate sloping lands, considering the
Menten parameter, was 24 weeks. Polyphenol content long-term conservation of land resource and the po-
in the biomass is usually higher and litter may be clas- tential for continuous cropping with minimal inputs. I.
siÞed as low-quality, but long durable mulch. Due to edulis has potential to restore degraded soils such as old

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pastures (TCA, 1997). There are examples of soil resto- FructiÞcation starts in the second year with few pods
ration in Latin America or in other countries such as in and increases till the Þfth year, when the production of
the Southern Cameroon (Kanmegne et al., 2003). one tree can reach as much as 300 pods (TCA, 1997).
I. edulis is particularly interesting to control the weed Mature fruits do not fall down the tree, but must be
development because, alone or in combination, this collected or shake-off by wooden sticks. The expected
species can have an integral shade cover. Alegre et al. time for production is about 20 year, but farmers typi-
(1999), for instance, have found signiÞcant effect of cally managed trees for fruit production for only 6–10
weed reduction the association between I. edulis and years, after which they are cut for Þrewood. Some farm-
Centrosema sp. ers coppice the trees for Þrewood in a two-year rotation
(ICRAF, 1996).
Silvicultural management
Integration into agroforestry systems
I. edulis is easily propagated by the seeds, which read-
ily germinate inside the pod. The fruits are indehiscent I. edulis is described as a tree which can be integrated
and embryos viviparous. The seeds cannot be stored in the agroforestry systems easily (Weber et al. 1997).
and must be planted immediately after opening the pot. The examples of this integration are particularly numer-
Germination starts since the third day and reaches the ous from Peruvian Amazonian, and also from several
rate of 90% (Villachica, 1996). Farmers usually selects other countries in Latin America. I. edulis and also other
the best pods and use direct sowding, making a small Inga species are most often associated with coffee or ca-
hole by the stick and putting 2–4 seeds inside. Concern- cao trees as shade tree (Brack, 1993).
ing the seeds selection, farmers prefer to sow black- The cultivation of I. edulis is also adapted to wide va-
coloured seeds of these varieties in their experience; riety of agroforestry association. It is possible to plant I.
black seeds develop into so called “female” trees with edulis with several annual, semi-perennial or perennial
abundant, annual fruit production while yellow seeds species. In alley-cropping, the potential associated crops
develop into “male” trees which do not produce much are cassava (Manihot esculenta), banana (Musa spp.),
fruit (Weber et al., 1997). The preparation of seedling in pineapple (Ananas comosus), caimito (Pouteria caimi-
the nursery and transplanting when they reach the height to), carambola (Averrhoa carambola), avocado (Persea
of 40–60 cm to the deÞnitive place is also possible. One americana), coconut (Cocos nucifera), citrus trees (Cit-
farmer in Ucayali region of Peruvian Amazon explained rus spp.), tropical cedar (Cedrela odorata), Brazilian nut
how he annually planted I. edulis in his crop Þeld with (Bertholletia excelsa), moena (Aniba sp.) and mahoga-
seed from his home garden, but also collected seed from ny (Swetenia macrophylla) (TCA, 1997). The practice
neighbours’ trees and from fruit bought in the market in of managing a fallow by small farmers in the Peruvian
a more haphazard rather than planned fashion. Selection Amazon actually begins at the time the forest is cleared
for speciÞed traits is often opportunistic (Brodie et al., for cultivation, when valuable species such as palms, and
1997). those used later for timber, are spared during cutting. As
For planting the species as fallow enrichment farmers the swidden is being cultivated, perennial species such
establish I. edulis in their annual crops by direct sowing as pineapple (Ananas comosus), banana (Musa spp.) and
at approximately 5 × 5 m density (Brodie et al., 1997). fruit and palm trees such as umarí (Poraqueiba sericea),
After the crop harvest, the Inga trees are allowed to grow uvilla (Pouroma cecropiaefolia) and peach palm (Bactris
and form a closed canopy. The farmer estimates that 2–3 gasipaes), among others I. edulis is planted, interspersed
years is needed to suppress weeds and restore the fer- among annual crops such as cassava (Manihot esculenta)
tility sufÞciently to be able to sow annual crops. How- or maize (Zea mays) (Unruch, 1988).
ever, farmers often decide to establish some fruit trees It can also be integrated in the crops succession. For
under the canopy of Inga trees, but they must be pruned example, near the city of Pucallpa (Peruvian Amazon),
or thinned to avoid excessive shade. As a shade trees for in the non-ßooding land, the annual crops are establish-
coffee and cacao the distance of 10–15 m among the trees ment for 1–3 years (rice, maize, cassava and other asso-
is used, while for hedgerow intercropping 4 m between ciated crops: beans, chickpea, squash, etc.) followed by
hedges and 0.5 m between the trees inside the hedge is 1–5 years of perennial crops, typically plantain (Musa
used (Villachica, 1996). There are also some experiences sp.), with associated fruit trees. As plantain yields de-
of establishment of pure plantation and it is expected to cline between the sixth and eighth years, they are sub-
maintain the distances of 3–6 m. The trees can reach the stituting plantains with fast producing fruit trees, most
height of 2–4.5 m in 12 months, depending on the soil commonly I. edulis, which are then succeeded by longer
fertility (Villachica, 1996). living marketable fruit trees such as citruses (Citrus spp.)

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and peach palm (Bactris gasipaes). These are often inter- its in the Þrst years, because of the higher establishment
cropped with pineapple, tree densities are low and man- and managements costs. It could take up to ten years for
agement extensive (Brodie et al., 1997). Other example smallholders to begin making a proÞt above that achiev-
of integration is 30-year multistrata systems (Weber et able with the traditional fallow system.
al., 1997), that could provide a diverse and sustainable Another study conducted at Yurimaguas by Fernandez
source of income to farmer: fruit and Þrewood from I. et al. (1993) included double hedgerows of I. edulis were
edulis beginning in year two; Þrewood, pulp and small established and an annual rotation of upland rice and
poles from bolaina (Guazuma crinita) and charcoal and cowpea provided the test crops over several years. The
small poles from capirona (Callycophylum sprucea- effects of prunings applied as a mulch, application of
num) beginning in year three; fruit, seed and “heart of fertilizers and repeated root pruning were investigated.
palm” from peach palm beginning in year Þve; lumber Mulching increased rice grain yields and reduced seed
and large poles for construction from bolaina and capi- yields of cowpeas. Recommendations included alley
rona beginning in year ten; and high-value lumber from widths >4m, intense hedgerow pruning, incorporation of
tornillo (Cedrelinga cateniformis) in year 30. The tree prunings into the soil, moderate fertilizer use, and use of
species could be sequentially interplanted in secondary species less prone to competition than rice. The species
forests, or planted separately in spatially distinct patches, is also used in the contour hedgerows.
strips, depending of the soil conditions. The home gardens are other example of agroforestry
Following the cropping cycle, many farmers in the systems where I. edulis is often planted. As Padoch and
Peruvian Amazon Basin plant Inga edulis into their Jong (1991) concluded from their study of species diver-
woody fallows (Brodie et al., 1997; Sotelo-Montes and sity of a riverside village near Iquitos (Peruvian Ama-
Weber, 1997). The use of I. edulis green manure for fal- zon), I. edulis was the most widely distributed tree spe-
low improvement is well developed in some idingenous cies. The tree is also used in the silvopastoral systems
farming communities: they can maintain sustainable in order to improve the soil physical and chemical char-
crop production for 5 years following a 5-year fallow acteristics (and thus the forage production) and also to
enriched with selected I. edulis (Brodie et al., 1997). In produce shade for cattle and marketable products. Weber
the trial of Alegre et al. (2005) I. edulis was used as a et al. (1997) described, for instance, the case of pastures
fallow tree. An experiment was conducted at Yurimaguas diversiÞed by planting timber species capirona (Calyco-
(Peruvian Amazon) evaluating planted tree fallows of I. phyllum spruceaneum) and I. edulis in low density or as
edulis and colubrina (Colubrina glandulosa), with and live fence around the pastures.
without centrosema (Centrosema macrocarpum) cover, In other regions of Latina America, most examples of
compared with the traditional bush fallow in terms of integration of I. edulis in the agroforestry systems can
weed suppression, their ability to increase subsequent also be found as shade tree for coffee or cacao (Brazil:
crop production and overall economic beneÞts. The Miranda et al., 1999; Costa Rica: Beer, 1991; Somarriba
planted trees grew faster and accumulated more biomass and Beer, 1999; Venezuela: Escalante et al., 1987). In
than those in natural fallow. The trees effectively sup- Brazil, Brienza et al. (1998) described a change of tradi-
pressed weeds, but the tree fallows did not increase crop tional slash-and-burn farming to improved fallow system
yields compared with natural fallow and resulted in sig- with the use of I. edulis, which was associated with the
niÞcantly lower yields in the third post-fallow season. maize and cassava. The tree biomass production was ac-
The Inga and Inga+centrosema systems had the highest celerated after cassava harvest. Cassava yield was not
net present value; however, natural fallow had the high- inßuenced negatively by tree growth, nor were the trees
est beneÞt/cost ratio, due to its lower costs relative to affected by the crop. Example of alley cropping in Brazil
the more intensive fallow systems. Long-term simulation is given by Ferraz et al. (2001): cowpeas and rice were
and economic assessment of the use of I. edulis as im- grown in rotation in alleys of I. edulis and other legumi-
proved fallow species compared to traditional bush fal- nous tree species, but cowpea seed yield did not respond
low in Peruvian Amazon (Lojka et al. 2008) has shown to the legume crop residues. Jardin et al. (2004) evalu-
that Inga fallow system can provide improvements to a ated the initial behaviour of a 32-month-old agroforestry
range of soil biophysical measures and enables higher system, where copoacú trees (Theobroma grandißorum)
levels of farm outputs to be achieved. The economic are intercropped simultaneously with forest species, I.
analysis has also demonstrated higher proÞtability of edulis used for permanent shading and with different va-
this system, but only in long term. This can be attributed rieties of banana (Musa spp.) for temporary shading. I.
to improved soil fertility (thus higher crop yields), and edulis presented a very rapid initial growth; however, it
to the additional product (Þrewood). But in adopting the may become inconvenient for use as a shade species, be-
Inga fallow system, smallholders will incur lower prof- cause of its horizontalized and very dense type of crown

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which produces excessive shading. Nonetheless, they be a potential species to be used. It is easily adapted into
conclude, that if employed with adequate spacing and agroforestry systems, it can be a good source of income
guided by pruning, it may be appropriate as a compo- by the fruit or wood sales for farmers and it is able to
nent of agroforestry systems, due to the large quantity of maintain and improve soil fertility. It is surprising that this
biomass produced, which functions to protect and enrich widely used tree species in Latin America is still not the
the soil. Miranda et al. (1999) studied the performance object of any improved breeding program and scientiÞc
of coffee plants shaded by I. edulis, compared to plants information about this species are still rather scarce.
exposed to full sun. The results of two harvests of coffee
showed that the yield in full sun was signiÞcantly higher.
Several examples of the use of I. edulis in agroforestry REFERENCES
systems take place in Costa Rica. Kettler (1997) stud-
ied the effects of enriched fallows systems with I. edulis ALEGRE J.C., FERNANDES E.C.M. (1991): Runoff and
and other leguminous tree species. Biomass production erosion losses under forest low-input and alley-crop-
of the fallow vegetation was of better quality and quan- ping on slopes: Y-433B. In: TropSoils Technical Re-
tity in all fallow enrichment treatments. However, bean port 1988–89. Raleigh, North Carolina State Univer-
yields did not show a signiÞcant response to the fallow sity, NC (USA), pp. 227–228.
enrichment treatments. In other study from Costa Rica, ALEGRE J.C., RAO M.R. (1995): Soil and water con-
timber species Terminalia amazonia was planted alone, servation by contour hedging in the humid tropics of
and interplanted with I. edulis (Nichols and Carpenter, Peru. Agr. Ecosyst. Environ., 57: 17–25.
2006). Measurements at 4, 8, and 11 years showed that ALEGRE J.C., AREVALO L., RAO M.R. (1999): Barbe-
T. amazonia grew signiÞcantly better when mixed with chos mejorados para un manejo intensivo de los sue-
I. edulis. They analyzed the standing crops of vegetation los en los trópicos húmedos. Seminario No 1 del Cen-
and their nitrogen content and the data were consistent tro Ecorregional, Pucallpa 15 de julio de 1999.
with the hypothesis that improved nitrogen nutrition ac- ALEGRE J.C., RAO M.R., AREVALO L.A., GUZMAN
counts for improved growth of T. amazonia when inter- W., FAMINOW M.D. (2005): Planted trees fallows for
planted with I. edulis. improving land productivity in the humid tropics of
A system being developed in Amazonian Ecuador is Peru. Agr. Ecosyst. Environ., 110: 104–117.
described (Bishop, 1983) in which the fallow period in BEER J. (1991): Implementing on-farm agroforestry
shifting cultivation sequences is intensiÞed by the use research: lessons learned in Talamanca, Costa Rica.
of the Asian perennial forage cover crop Desmodium Special issue: On-farm agroforestry research, based
ovalifolium under I. edulis. Both species improve soil on an international workshop “Methods for partici-
fertility and reduce erosion. I. edulis produced fuelwood patory on-farm agroforestry research” Nairobi, 1990.
and charcoal in less than six years, produced fruit and Agroforestry Syst., 15: 229–243.
provided good bee forage. The system was compatible BISHOP J.P. (1983): Tropical forest sheep on legume fo-
with traditional patterns of cultivation, forming a six- rage/fuelwood fallows. Agroforestry Syst., 1: 79–84.
year fallow in an eight-year Þeld rotation with maize, BRACK E.A. (1993): Plantas nativas utilizadas en el Perú
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SZOTT L.T. (1987): Improving the productivity of shift- Received for publication on March 26, 2010
ing cultivation in the Amazon basin of Peru through the Accepted for publication on October 18, 2010

Corresponding author:
Ing. Bohdan Lojka, PhD.
Institute of Tropics and Subtropics
Czech University of Life Sciences Prague
Kamýcká 129
165 21 Prague 6-Suchdol
Czech Republic
email: lojka@its.czu.cz

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