Plant and Soil 222: 59–70, 2000.

© 2000 Kluwer Academic Publishers. Printed in the Netherlands.

59

Land quality changes following the conversion of the natural vegetation

into silvo-pastoral systems in semi-arid NE Brazil

Barbara Wick1,∗ , Holm Tiessen2 and Romulo S.C. Menezes3

1 Institute
of Soil Science and Forest Nutrition, Georg August University Göttingen, Büsgenweg 2, D-37077 Göttin-
gen, Germany; 2 Department of Soil Science, University of Saskatchewan, 51 Campus Drive, Saskatoon, Canada
S7N 5A8 and 3 Natural Resource Ecology Laboratory, Colorado State University, Fort Collins, CO 80521, USA

Received 18 August 1999. Accepted in revised form 2 February 2000

Key words: indigenous trees, land quality change, microbial biomass, nutrient dynamics, silvo-pastoral systems,
soil enzymes

Abstract
The presence of trees in grasslands frequently results in resource islands of higher soil quality. Therefore, some nat-
ive trees are often preserved or agroforestry species are interplanted when land is cleared for pasture. We evaluated
soil quality changes associated with the conversion of a native thorn forest (caatinga) into silvo-pastoral systems
in semiarid NE Brazil. Soil nutrients, organic matter, microbial biomass and soil enzymes under native caatinga,
the canopy of two preserved native (Ziziphus joazeiro and Spondias tuberosa) and one introduced tree species
(Prosopis juliflora), and under a planted pasture of Cenchrus ciliaris were compared. The two preserved tree
species maintained high nutrient and organic matter contents and high biological activity levels not only relative
to the grass but also relative to the native caatinga. The non-deciduous Z. joazeiro was more effective than the
deciduous S. tuberosa, and enriched surface soils by 100 mg P kg−1 . The complete replacement of natural caatinga
with C. ciliaris pasture or a C. ciliaris–P. juliflora silvopastoral system had, after 14 years, decreased microbial
biomass C and N contents, and ß-glucosidase activity, but did not affect soil nutrient or organic matter status.
The biological activity under the trees of the tree-grass association was greater than under grass alone, although
P. juliflora did not maintain higher nutrient or organic matter levels. The search for indigenous alternatives to the
main agroforestry species may provide viable alternatives for improved land management and the conservation of
biodiversity. Microbiological and biochemical parameters responded more readily to changes in land management
than chemical fertility indices. They are important indices for the impact of vegetation changes and its associated
land quality changes.

Introduction Natural forests maintained or improved soil fertil-

Changes in land cover associated with different land
uses are important agents of environmental change
and degradation in the semiarid tropics. In the in-
terior of NE Brazil, the main land use change is the
conversion of natural vegetation to arable or grazing
lands. The slash and burn of the natural thorn forest
(caatinga) vegetation has led to a decline in land qual-
ity through organic matter and nutrient depletion and
erosion (Nunes Filho et al., 1990; Tiessen et al., 1992).
∗ FAX No: +49551 393310. E-mail: bwick1@gwdg.de
ity by closed nutrient cycling (Rhoades, 1997; Young,
1997). In semiarid pastoral ecosystems, trees are re-
cognized to improve soil conditions and herbaceous
productivity (Belsky, 1994; Rhoades, 1997). Soils
under tree canopies were reported to have higher
concentrations of organic matter and increased nu-
trient availability compared with neighboring grass-
lands (Garcia-Miragaya et al., 1994; Tiedemann and
Klemmedson, 1973). Shrubs and trees in arid and
semiarid environments also enhanced the abundance
and activity of the microbial biomass (Kieft, 1994).
Trees improved micro-climatic conditions under the
60
Table 1. Comparison of soil chemical characteristics under buffel grass pasture at different
locations within the beef cattle ranch

BJ a BU BC1 BC2 BC3 BC4 BA LSD b

(p=0.05)

pH 6.9 6.7 6.7 6.8 7.2 7.2 7.1 0.4

Ctot (g kg−1 ) 9.6 11.7 11.5 9.2 14.9 18.5 20.7 4.9
Ntot (g kg−1 ) 0.86 1.0 0.93 0.72 1.23 1.50 1.8 0.3
C4-C (g kg−1 )c 3.4 3.0 2.8 1.7 5.6 6.8 7.1 1.6
Ca (cmol+ kg−1 ) 3.8 4.4 3.9 2.6 6.9 8.8 8.2 2.5
Mg (cmol+ kg−1 ) 1.0 1.25 0.9 0.57 1.84 1.93 2.69 0.6
K (cmol+ kg−1 ) 0.68 0.64 0.51 0.35 1.05 1.02 1.16 0.3
Ptot (mg kg−1 ) 219 206 140 165 214 325 418 56
resin-Pi (mg kg−1 ) 14.6 7.3 3.3 3.6 5.8 6.5 7.9 4.5
Po (mg kg−1 ) 33 49 45 36 58 84 120 16
a B = buffel surrounding joazeiro; B = buffel surrounding umbuzeiro; B
J U C1 = buffel along
caatinga 1; BC2 = buffel along caatinga 2; BC3 = buffel along caatinga 3; BA = buffel sur-
rounding algaroba; BC4 = buffel along caatinga 4 surrounding algaroba trees AC4 .
b The number of comparisons were n = 8.
c C4-C is carbon derived from C-4 metabolic plants, i.e. principally the buffel grass that
replaced the predominantly C-3 caatinga.

canopy through the accumulation and retention of soil
moisture, reduced evaporation and lower temperature
(Knoop and Walker, 1985; Weltzin and Coughen-
our, 1990). The co-existence of savanna trees and
below-crown grasses have been explained by a two-
layer-model that delineates soil water uptake zones
spatially. Trees have access to deeper water stores
while grasses access moisture held in surface soils
(Knoop and Walker, 1985; Sala et al., 1989). Trees
can also increase understorey herbaceous productiv-
ity over that of open grassland (Belsky et al., 1993;
Weltzin and Coughenour, 1990) and provide forage for
browsing animals which, in turn, introduce nutrients
through manure beneath the canopy. Higher germin-
ation rates in the accumulated litter layers have been
reported (Kellman, 1979). The small-scale enriched
zones below trees and shrubs in arid and semi-arid
environments have been described as islands of fer-
tility or resource islands. They have been recorded
in savanna ecosystems in Africa (Belsky et al., 1993;
Weltzin and Coughenour, 1990) and central and south
America (Garcia-Miragaya, 1994; Kellman, 1979;
Menezes et al. 1999).
The positive effects of trees in restoring soil quality
are employed in the tree fallows of shifting cultiv-
ation systems, and in agroforestry or silvo-pastoral
land use systems that maintain a permanent associ-
ation between trees and crops or pastures. Some land
use systems that employ tree associations are tradi-
tional (home gardens, living fences or hedges, grazing
parklands) others are being developed in efforts to find
agro-silvo-pastoral alternatives to slash-burn landuses.
The range of trees tested in new silvo-pastoral systems
is often restricted, but recent initiatives increasingly
aim to maintain biodiversity. The domestication of in-
digenous forest trees has been proposed as a means of
intensifying and diversifying land use in a way that is
both sustainable and productive (Sanchez et al., 1997),
and integration of trees into tropical farming systems
is being promoted (Leaky and Simons, 1998).
Many agroforestry initiatives have involved the in-
troduction of fast growing and (potentially) N-fixing
exotic trees. Algaroba (Prosopis juliflora) has been
introduced to NE Brazil in the 1980s because it was
considered superior to native trees (Gibson, 1992).
More traditionally, farmers and ranchers preserve se-
lected native trees on agricultural and grazing lands.
The species selection is based on such benefits as
shade, fruit or wood quality. Two trees commonly
preserved in semiarid NE Brazil are the umbuzeiro
(Spondias tuberosa), valued for its fruit and shade, and
the joazeiro (Ziziphus joazeiro) which provides shade
and browse material.
The present study was undertaken to evaluate the
soil quality associated with these introduced or pre-
served trees in a silvo-pastoral system. Soil nutrients,
organic matter and microbiological characteristics un-
der the canopy of two preserved native and one planted
exotic tree species were compared with those found
 61
Table 2. Impact of caatinga clearing and establishment of buffel grass pasture within preserved caatinga trees on soil chemical properties

Land cover / Test Total C Total N Resin-P Po Total P Ca Mg K pH

comparison (g kg−1 ) (g kg−1 ) (mg kg−1 ) (mg kg−1 ) (mg kg−1 ) (cmol+ kg−1 ) (cmol+ kg−1 ) (cmol+ kg−1 )

caatinga (C1,C2,C3) 14.1 1.1 4.2 52 176 5.0 1.1 0.42 6.7
buffel (BC1, BC2, BC3 ) 12.0 0.98 4.2 46 173 4.5 1.1 0.64 6.9
joazeiro (J) 18.4 1.61 32.3 58 308 6.6 1.6 1.04 7.3
single joazeiro in C1 17.4 1.16 3.6 48 144 5.1 1.4 0.3 6.3
umbuzeiro (U) 15.4 1.3 10.8 65 239 5.1 1.32 1.00 7.0
single umbuzeiro in C1 20.4 1.4 6.0 89 231 6.3 1.5 0.6 6.3

caatinga / buffel ta ns ns ns ns ns ns ns 0.22 ∗∗ 0.2∗

(C1,2,3 / BC1, BC2, BC3 )
caatinga / joazeiro LSDb 3.6 0.3 7.8 ns 31 1.4 0.3 0.16 0.2
(C1 / C2 / C3 / J)
caatinga / umbuzeiro LSDb ns ns 2.5 9.8 32 ns ns 0.25 0.23
(C1 / C2 / C3 / U)
a Significance level of paired comparison (t-test) of caatinga versus buffel grass was: ∗ at p < 0.05; ∗∗ at p < 0.01; ∗∗∗ at p < 0.001; the number
of paired comparisons were n=24. Values are the difference between the soils.
b = 0.05.
ns = non significant.

outside the canopy in the planted pasture and those
under native intact thorn forest (caatinga).
Materials and methods
Study region
The sampling sites were located in a beef cattle
ranch at Custodia, Pernambuco, NE Brazil (8◦ 140
S, 37◦ 450 W). The native vegetation was caatinga,
a semi-deciduous thorn forest (Mors, 1994), which
is traditionally used for fuelwood, and slash-and-
burn shifting cultivation followed by extensive graz-
ing and browsing (Sampaio, 1995). Rainfall averages
about 600 mm y−1 , concentrated between January and
May with a nearly rainless period between June and
December (Sampaio, 1995). Rainfall is highly erratic
and extended droughts occur once or twice every dec-
ade (Eiten, 1992). Mean annual temperature is 26 ◦ C.
The soils are non-calcic brown (Ustalfs) on crystalline
rock parent material with an average sand content of
75%.
Sampling sites
Until 1984, the study area was covered with native
caatinga at least 50 years old. In 1984, 3000 ha were
completely cleared with chains, and buffel grass (Cen-
chrus ciliaris L., a perennial tufted bunchgrass native
to southern Africa, Meredith, 1955, as quoted by
Wilson, 1996) was planted. One year later, algaroba
(Prosopis juliflora, Swartz D.C.) was interseeded in
the buffel grass pasture at a spacing of 10 × 10 m.
Algaroba is an evergreen, potentially N-fixing tree nat-
ive to central America (Babos and Cumana, 1992). Its
wood is used for fuel and construction, and the pods
and leaves as fodder. The gum exuded from the sap-
wood is used in the paper and cosmetic industry (De
Oliveira Antonino, 1992). At the time of sampling in
1998, the tree plantation was 13 years old.
On the remaining area of the farm (> 3000 ha)
the land clearing of 1984 preserved two native caat-
inga species. Joazeiro (Ziziphus joazeiro Mart.) is a
non-deciduous tree used for medicinals, fodder and
fuelwood; its fruits are edible but not marketable
(Noel, 1995). Umbuzeiro (Spondias tuberosa Arruda
Cam.) is deciduous and produces large quantities of
marketable fruit. The wood is used for construction
(Machado et al., 1997; Mors, 1994). The preserved
mature trees were spaced some 30–40 m apart in the
14-year old continuous pasture. Caatinga regrowth in
the buffel grass and around the algarobas was slashed
every other year.
Soil sampling and processing
Top soil (8 cm) samples were taken in February 1998.
A total of 16 algarobas were sampled from two sites
about 700 m apart. Paired samples were taken for
 62

each tree – one sample from under the canopy and

one 5 m away from the canopy edge in the buffel

a Significance level of paired comparison (t-test) of caatinga versus buffel grass was: ∗ at p < 0.05; ∗∗ at p < 0.01; ∗∗∗ at p < 0.001;. the number of paired comparisons were n=24.
grass pasture. The first site within the plantation (eight
Table 3. Impact of caatinga clearing and establishment of buffel grass pasture within preserved caatinga trees on soil microbiological properties and light fraction (LF) N and P

(µg tyrosine
g−1 2h−1 )
Protease
trees, samples A from under the canopy and BA from

8.4
22

26
42

13
27

61

71
ns
the paired buffel location) was a clean stand of buffel
grass interspersed with algaroba. The trees at this site
were chosen randomly. The second site (eight trees,
(µg PN g−1 h−1 )
glucocosidase

samples AC4) was adjacent to an undisturbed caatinga

22 ∗∗∗
136
187
42

34

25
64

49

87
ß-

site (C4). Here, eight trees (AC4 ) were chosen in a

transect parallel to the edge of the caatinga. The as-
sociated buffel grass (samples BC4 ) was not as clean
as that at site A, and invading shrubs had recently
(µg PN g−1 h−1 )
phosphatase

been slashed. Within the caatinga (C4; n=8) samples

Alkaline

124

199
166

195
140
384

were also taken along a transect parallel to the one of

82
ns

ns
AC4 and its associated buffel samples (BC4 ) so that
spatially paired samples were obtained between the
caatinga and the adjacent cleared site.
(µg PN g−1 h−1 )

In the area where clearing had been selective, eight

phosphatase

preserved trees each of joazeiro and umbuzeiro were

Acid

264

403
335

598
412
389

ns

ns

ns

randomly chosen and sampled in the following way:

three top soil samples were taken from under the
canopy of the joazeiros (J) and umbuzeiros (U). The
(mg kg−1 )

distance of each sample to the tree bole as well as

Mic-P

1.4 ∗
3.8

1.9
5.6
6.3
5.2

8.0
4.0

ns

the direction was selected at random. Corresponding

samples paired to each of the under-canopy samples
were taken from the buffel grass pasture (BJ , BU )
(mg kg−1 )

following the same direction away from the bole 5 m

Mic-N

6.3
4∗
16

21
25
20

23
31

ns

beyond the canopy edge.

Three undisturbed caatinga sites (C1, C2, C3) were
also sampled at eight random spots each along tran-
(mg kg−1 )
Mic-C

sects parallel to the forest edge. Paired sampling was

131
135
107

167
83

29
76

ns

ns

then done about 10 m outside the caatinga in adjacent

buffel grass sites (BC1 , BC2 , BC3 ).
(mg kg−1 )

As a result of this sampling scheme, we obtained

LF-Po

20.5
21.9
16.5

18.7
28.9
nd

nd

10

ns

samples for pairwise comparisons of each tree spe-

cies with surrounding buffel of caatinga with adjacent
buffel, and of caatinga with adjacent algaroba. In
(g kg−1 )

addition, group comparisons were possible between

LF-N

0.3
0.4
0.4
0.4

0.6
0.8
nd

nd

Values are the difference between the soils.

ns

ns = non significant; nd = not determined.

four caatingas, the three tree species and five areas of

buffel.
LSDb

LSDb

The soil cores were taken with an auger of 8.25 cm

Test

ta

diameter, transferred into sealed plastic bags and

transported in styrofoam coolers to the laboratory
(C1,2,3 / BC1 ,BC2 ,BC3 )
single umbuzeiro in C1
buffel (BC1 ,BC2 ,BC3 )

within 3 days. Subsamples for the determination of

caatinga / umbuzeiro
single joazeiro in C1
caatinga (C1,C2,C3)

caatinga / joazeiro

(C1 / C2 / C3 / U)
(C1 / C2 / C3 / J)
caatinga / buffel
umbuzeiro (U)
Land cover /

pH, cations and water-floatable organic matter N and

comparison

joazeiro (J)

P (light fraction) were air-dried and sieved through

a 2 mm sieve. Subsamples for total C, N and P, and
b = 0.05.

sequential fractionation of phosphorus were air-dried

and ground to < 150 µm on a ballmill. The sub-
samples for enzyme analysis and microbial biomass
 63
Table 4. Average soil chemical characteristics under joazeiro and umbuzeiro trees left following caatinga clearing versus open buffel grass

Land cover / Test total C Total N Resin-P Po Total P Ca Mg K pH

comparison (g kg−1 ) (g kg−1 ) (mg kg−1 ) (mg kg−1 ) (mg kg−1 ) (cmol+ kg−1 ) (cmol+ kg−1 ) (cmol+ kg−1 )

Joazeiro (J) 18.4 1.6 32.3 58 308 6.6 1.6 1.04 7.3
Buffel (BJ ) 9.7 0.86 14.6 33 219 3.8 1.0 0.69 6.9
Umbuzeiro (U) 15.4 1.3 10.8 65 239 5.1 1.32 1.00 7.0
Buffel (BU ) 11.7 1.0 7.3 49 206 4.4 1.25 0.64 6.7

J / BJ ta 8.7 ∗∗∗ 0.7 ∗∗∗ 18 ∗∗∗ 25 ∗∗∗ 90 ∗∗∗ 2.8 ∗∗∗ 0.6 ∗∗∗ 0.35 ∗∗∗ 0.4 ∗∗∗
U / BU LSDb 3.7 ∗∗ 0.3 ∗∗ 3.5 ∗ 16 ∗∗∗ 33 ∗ ns ns 0.35 ∗ 0.3 ∗∗
J / U / BJ / BU LSDb 3.5 0.26 8.6 10 52 1.1 0.32 0.25 0.3
a Significance level of paired comparison (t-test) of tree versus buffel grass was: ∗ at p < 0.05; ∗∗ at p < 0.01; ∗∗∗ at p < 0.001. The number of
paired comparisons were n=24. Values are the difference between the soils.
b = 0.05.
ns = non significant.

determination were sieved to pass 2 mm and were kept
field moist at 4 ◦ C until analysis.
Soil analysis
Total C and N (and 13 C and 15 N abundance, de-
tails not shown) were measured on a continuous flow
isotope ratio mass spectrometer (Europa scientific).
Total P was analyzed by a H2 O2 /H2 SO4 digestion and
autoanalysis (Thomas et al., 1967). Available P was
determined by resin extraction, and organic P (Po)
by summation of hydroxide and hot concentrated acid
extractable organic P fractions of an sequential ex-
traction simplified by omitting the bicarbonate extract
from Tiessen and Moir (1993).
Calcium, Mg and K were extracted by unbuffered 1
M NH4 Cl. Calcium and Mg were analyzed by atomic
absorption, and K by emission spectrophotometry; pH
was measured in a 1:10 soil:water suspension.
The light organic matter fraction was separated by
sonifying a soil:water suspension of 20 g soil and 150
ml deionised water for 3 min at 300 W output with a
Braun-sonic sonifier. During sonification, the suspen-
sion was cooled in an ice-water bath. The suspension
was passed through a 300 mesh sieve to retain both
the sand and the water-floatable light organic matter
fraction. The retained fraction was mixed and ground
in a mortar after oven drying at 60 ◦ C. Organic P in the
light fraction was determined according to the method
of Saunders and Williams (1955).
Soil microbiological analysis
Acid and alkaline phosphatase, and ß-glucosidase
activity in the soil were measured according to
Tabatabai and Bremner (1969) and Eivazi and
Tabatabai (1988), respectively. Toluene was not in-
cluded in the procedure. Results of activity were
expressed as µg p-nitrophenol released over 1 h by 1 g
dry soil and are averages of triplicate assays.
Protease (casein-hydrolyzing) activity of the soil
was determined according to Ladd and Butler (1972).
Results of activity are expressed as µg tyrosine re-
leased over 2 h by 1 g dry soil. Microbial biomass C
and N were determined by the chloroform-fumigation-
extraction method of Vance et al. (1987). The soil
was adjusted to 30% water holding capacity (WHC)
prior to fumigation and extraction. Total organic C of
the K2 SO4 -extracts was analyzed by a chromic acid
digestion, and the Cr3+ -product measured spectropho-
tometrically (Heanes, 1984). Total N of the K2 SO4 -
extracts were determined by persulphate oxidation
according to the method of Raveh and Avnimelech
(1979). The released ammonium was measured on a
Technicon II Autoanalyzer. Microbial biomass C and
N were calculated by subtracting the amount extracted
from the unfumigated soils from that extracted from
the fumigated soils.
Microbial biomass P analysis was adapted from
Kouno et al. (1995). Inorganic phosphorus of the bio-
mass released by chloroform fumigation for 24 h was
extracted with resin strips and determined according to
the method of Tiessen and Moir (1993) on a Technicon
II Autoanalyzer. Biomass P was calculated as P ex-
tracted by the resin strips after 24 h fumigation less the
amount extracted by the anion strips from unfumigated
soils.
64

Statistics

Statistical analysis was performed with the Systat-

Program (SYSTAT, 1992) by one-factorial ANOVA
using land cover (caatinga, trees, grass) as levels of
Table 5. Average soil microbiological and light fraction (LF) N and P characteristics under joazeiro and umbuzeiro trees left following caatinga clearing versus

∗ Significance level of paired comparison (t-test) of tree versus buffel grass was: ∗ at p < 0.05; ∗∗ at p < 0.01; ∗∗∗ at p < 0.001; the number of paired comparisons
(µg tyrosine
(g kg−1 ) (mg kg−1 ) (mg kg−1 ) (mg kg−1 ) (mg kg−1 ) (µg PN g−1 h−1 ) (µg PN g−1 h−1 ) (µg PN g−1 h−1 ) g−1 2 h−1 )
Protease

one factor. Differences under the tree canopies versus

37 ∗∗∗
15 ∗∗
61

42
24

27

12
outside the canopies in the grass were determined
by the dependent t-test for pairwise comparison. The
number of paired comparisons for joazeiro and um-
buzeiro versus buffel grass were 24 each, and for
glucosidase

115 ∗∗∗
67 ∗∗∗
algaroba versus buffel grass 16. A pairwise compar-
187

136
72

69

39
ß-

ison was also performed between caatinga 1 and 3 and

the corresponding buffel grass (BC1 , BC2 , BC3 ). The
number of paired comparisons were 24.
phosphatase
Alkaline

256 ∗∗∗
74 ∗∗
384

199
128

125

69

Results and discussion

Soil spatial variability of the sampling sites

To determine vegetation effects on soil, we needed

phosphatase

189 ∗∗∗
108 ∗

to evaluate the soils spatial variability, since both the

Acid

389

403
200

295

116

were n=24 for total soil and n=8 for the light fraction. Values are the difference between the soils.

presence of specific trees and their influence on soil

conditions may depend on original soil properties at
both microsite and landscape scales (Rhoades, 1997).
It is, therefore, possible that measured soil properties
4.4 ∗∗∗
Mic-P

2.2 ∗∗
8.0

5.6
3.6

3.4

1.5

are not only produced by the tree but that the pres-
ence of the tree is a result of the soil properties. We,
therefore, examined soils under identical vegetation
Mic-N

17 ∗∗∗
7 ∗∗

covers, i.e. the buffel grass planted on cleared caatinga

5.5
31

21
14

14

14 years prior to our sampling, and also the four sites

of remaining caatinga. Average soil chemical charac-
Mic-C

69 ∗∗∗

teristics at the different buffel grass locations (Table 1)

27 ∗
167

131
104
98

30

show that both BA and BC4 had the highest soil nutri-
ent status and grass productivity. These two sites (and
the adjacent areas discussed below) were consequently
19.6 ∗∗
LF-Po

7.3 ∗
28.9

20.5
13.2

10.3
9.3

treated separately in the data analysis.

With the exception of available phosphorus, all
properties measured under BJ were not statistically
0.55 ∗∗
0.18 ∗
LF-N

0.78

0.44
0.23

0.26

0.29

different from BU , BC1 and BC2 . The soils most likely

to be different in this landscape might have been those
surrounding joazeiros (BJ ) since in the more arid parts
LSDb
J / U / BJ / BU LSDb
Test

ta

of the NE, these trees are often associated with the

ns = non significant.
open buffel grass

lowest landscape positions that may be covered by

Umbuzeiro (U)
Land cover /

Buffel (BU )
Buffel (BJ )
Joazeiro (J)
comparison

alluvial materials from surrounding reliefs (Machado

U / BU
J / BJ

et al., 1997). The absence of significant differences

b =0.05.

between BJ and the other buffel sites indicates that

the joazeiro are growing on soils similar to those of the
other sites, and that the tree effects can be examined.
BC3 had somewhat higher soil organic matter
(SOM) and exchangeable cations than BJ , BC1 and
BC2 but was not statistically different from BU . The

higher SOM and base status could be attributed to
the higher grass productivity since C contents with a
13 C signature of C-4 plants (i.e. buffel grass) at B
C3
were significantly higher than at BJ , BU , BC1 and
BC2 (Table 1). Carbon derived from the trees (C3-C)
was not significantly different between the sites (data
not shown). Therefore, the soil under BC3 was not
considered different from BJ , BU , BC1 and BC2 .
The comparison of the four caatinga sites (C1–
C4) showed the same pattern as the comparison of the
buffel grass sites BC1 –BC4 with slightly lower carbon
and exchangeable bases in C2 and elevated phosphorus
in C4 (data not shown). Based on these results we dis-
tinguished 2 areas on the ranch that had significantly
different soil properties. The first and larger area con-
tained all sites except those of A, BA , C4, AC4 and
BC4 , which together constituted the second area.
Selective clearing of caatinga
Buffel grass
The replacement of the caatinga with a buffel grass
pasture has, after 14 years, resulted in 50% higher ex-
changeable K in the pasture and a pH rise from 6.7 to
6.9 (Table 2). No impoverishment of the soil nutrient
or organic matter status was seen. Literature on forest
to pasture conversion reports conflicting results. For
sites in Mexico (Garcia-Oliva et al., 1994, 1999) and
Costa Rica (Veldkamp, 1994), net losses of organic
matter were reported while Buschbacher et al. (1988)
reported no change and Cerri et al. (1991) reported C
gains in Amazonian pastures.
Microbial biomass P and N as well as ß-
glucosidase (Table 3) were reduced under the grass
relative to the caatinga by 30%, 20% and 35%, re-
spectively. Microbial biomass and ß-glucosidase are
closely associated with SOM turnover, and both may
respond more readily than SOM as a whole to changes
in management (Powlson, 1994; Wick et al., 1998).
Conservation of caatinga trees
Relative to the mean values in caatinga soils, the soil
under the joazeiros preserved during caatinga clear-
ing was higher in all measured parameters except
organic P (Table 2). This difference was particularly
pronounced for the soil microbiological parameters
(Table 3). Microbial biomass C and ß-glucosidase
activity were enhanced under joazeiro by 60% and
190%, respectively. Total C was 30% higher than the
caatinga mean but LF-C was not different (data not
shown).
65
Available P (660%), total P (75%) and alkaline
phosphatase activity (130%) were much higher under
the preserved joazeiros than under caatinga. Organic
P was not different. Since total P is usually determ-
ined by soil parent material, and only modified slowly
by pedogenesis and biocycling, an increase of total P
of some 100 mg kg−1 requires further examination.
We therefore compared the values under the preserved
joazeiros with those under the only joazeiro we had
found in the natural caatinga (Tables 2 and 3). This
comparison, although not statistically sound, showed
that most parameters were much higher under the isol-
ated trees, i.e. the joazeiro in the caatinga does not
show the same enrichment as the isolated joazeiros in
the pasture. However, it is possible that this joazeiro
was affected by localized runoff, and sand content
in the soil was 10% greater than under the isolated
joazeiros or the means of the caatingas. Since no caat-
inga was left at any lower slope positions, we could
not compare appropriate joazeiros at the same land-
scape positions. As seen below, comparisons between
joazeiro and associated buffel soils confirm that the
increased concentrations may indeed be a tree effect.
Several explanations might be advanced for the greater
nutrient contents under isolated joazeiros: 1. sheltering
animals may have brought in nutrients. Against this
is the evidence from 13 C values (not shown) that in-
dicate no enrichment in C4-C (the main food source
of grazers) under isolated joazeiros. 2. The access to
nutrients and rooting patterns may be different at the
lower slope sites. In this case, the enrichment of SOM
and nutrients under the solitary joazeiros would be the
result of an interaction between tree effect and land-
scape position, i.e. specific to the position of the tree
on alluvial material with greater moisture availability,
different rooting patterns or possibly the accumula-
tion of eroded materials under the tree. Menezes et
al. (1999), using a different sampling pattern and stat-
istical approach in the same landscape, also reported
considerably higher nutrient levels under joazeiro than
in surrounding areas.
Soils under the preserved umbuzeiros were signi-
ficantly higher than the mean caatinga values only for
P pools, exchangeable K (Table 2) , ß-glucosidase
and protease activities (Table 3). Comparison of an
umbuzeiro inside the caatinga with the isolated trees
showed no obvious difference (Tables 2 and 3).
Differences between preserved trees and buffel grass
The difference in soil properties under the pre-
served trees versus buffel grass pasture was evaluated
66
Table 6. Average soil chemical characteristics under caatinga, algaroba and buffel grass

Land cover / Test Total C Total-N Resin-P Po Total P Ca Mg K pH

comparison (g kg−1 ) (g kg−1 ) (mg kg−1 ) (mg kg−1 ) (mg kg−1 ) (cmol+ kg−1 ) (cmol+ kg−1 ) (cmol+ kg−1 )

Caatinga (C4) 16.5 1.5 4.7 113 326 7.1 1.9 0.6 6.8
Algaroba (A) 17.6 1.7 10.3 110 417 9.4 2.8 1.23 7.1
Buffel (BA ) 20.7 1.8 7.9 120 417 8.2 2.7 1.16 7.1

A / BA ta ns ns ns ns ns ns ns ns ns
C4 / A / BA LSDb ns ns 3.9 ns ns ns ns 0.12 ns
a Significance level of paired comparison (t-test) of tree versus buffel grass was: ∗ at p < 0.05; ∗∗ at p < 0.01; ∗∗∗ at p < 0.001. The
number of paired comparisons were n=8.
b =0.05.
ns = non significant.

by comparing values under the canopy with paired
samples outside the canopies. Total C and N (Table 4)
as well as microbial biomass C and N (Table 5) were
some 90% higher under the tree canopies. The en-
hancement for the LF-C (170%; data not shown) and
LF-N (240%) was even higher. Protease activity under
the trees was 150% and ß-glucosidase activity 160%
higher than in the buffel grass (Table 5). Higher micro-
bial biomass values had similarly been reported under
Ziziphus mucronata (native to the semi-arid savanna in
Zimbabwe) (Campbell et al., 1994).
Total P was 40% higher under the tree, while or-
ganic P was enhanced by 80% and available P by
120% (Table 4). The latter is reflected in greater mi-
crobial P (130%) and alkaline phosphatase activity
(200%), but also in enhanced LF-Po (210%; Table 5).
Increased levels of available P were also reported for
the much smaller Ziziphus spina-christi native to semi-
arid NE Nigeria (Verinumbe, 1993). The extensive
root system of joazeiro (Sampaio, pers. comm.) may
have contributed to nutrient uptake and accumulation
under the canopy (Young, 1997).
Can the higher total P under joazeiro be ex-
plained by nutrient import and recycling? The excess
P amounts to approximately 100 mg kg−1 . The area
under the canopy of an average joazeiro was near 50
m2 . The soil on this area to 8 cm depth will weigh
some 5 Mg. The excess P in that soil weight amounts
to 500 g total P. We measured a P concentration of
1.4 mg g−1 in joazeiro leaves, and estimate annual
litter production at 6 Mg ha−1 yr−1 (Tiessen et al.,
1998), or 30 kg per canopy. The increase in total P can
be explained if one third of the litter fall over the min-
imum age of the tree (50 yr) represents a net import by
deeper or more extensive roots.
The amounts of exchangeable basic cations were
much higher under the tree canopies than under buffel
grass. Assuming an average CEC of clay (kaolinite)
of 5 cmol+ kg−1 and of OM of 200 cmol+ kg−1 the
higher clay and OM contents under the tree would be
expected to contribute 3.3 cmol+ kg−1 . This is less
than the measured difference between trees and grass
(Table 4), i.e. over and above the CEC increase there
is an enrichment with bases, also shown in an increase
in pH under the canopies.
All soil microbiological parameters under joazeiro
were significantly elevated over those of the paired
pasture and the umbuzeiro sites (Tables 5). Total and
available P were increased the most (Table 4). But total
N, LF-N and Ca were also significantly higher. The
90 mg kg−1 higher total P under the canopy compared
with the surrounding pasture, determined in a pairwise
comparison of samples that were no more than 10 m
apart indicates that improved soil fertility and nutrient
cycling under joazeiro is likely to be a true effect of
the tree and its interactions with the soil.
The paired comparison of the soil chemical prop-
erties (Table 4) between under- and outside-canopy
positions for the umbuzeiro showed 124–165% higher
values for most parameters (except Ca and Mg). Light
fraction N and P, and all microbiological parameters
(Table 5) were significantly higher under the trees than
in the buffel grass.
The difference in nutrient pools under umbuzeiro
was much less than that under joazeiro. This is in
accordance with the comparison between both trees
and the caatinga (Tables 2 and 3). Relative to um-
buzeiro total N, exchangeable Ca and total P were
120% higher, microbial biomass and light fraction N
and P were 40–80% higher and enzyme activities were
40–100% higher under joazeiro. Evergreen trees (like
 67

the joazeiro) from a seasonal savanna in north cen-

tral Venezuela were similarly reported to have higher
SOM, K and ECEC values than deciduous ones. In
that study, the enhanced fertility under the evergreens
was attributed to the slower leaf decomposition rates

a Significance level of paired comparison (t-test) of tree versus buffel grass was: ∗ at p < 0.05; ∗∗∗ at p < 0.01; ∗∗∗ at p < 0.001. The number of paired
(µ tyrosine
(g kg−1 ) (mg kg−1 ) (mg kg−1 ) (mg kg−1 ) (mg kg−1 ) (µg PN g−1 h−1 ) (µg PN g−1 h−1 ) (µg PN g−1 h−1 ) g−1 2 h−1 )
Protease
and to different N and P tissue concentration (Garcia-

51
49
40

ns
ns
Miragaya et al., 1994). However, we did not find any
significant differences of N (21 and 19.8 mg g−1 for
joazeiro and umbuzeiro, respectively) or P (1.4 and
1.5 mg g−1 for joazeiro and umbuzeiro, respectively)
glucosidase

26 ∗∗
83

16
89
63
in the leaf tissue.
ß-

Complete replacement of caatinga with algaroba and

Table 7. Average soil microbiological characteristics and light fraction (LF) N and P under caatinga, algaroba and buffel grass

buffel grass plantation

phosphatase
Alkaline

35 ∗∗
246
336
271

The elimination of the natural caatinga and replace-

ns

ment by buffel grass pasture for 14 years did not alter

the chemical (Table 6) and microbiological properties
(Table 7) of the soil. The only differences seen were
an increase of K (90%) and a decrease of microbial
phosphatase

biomass N and ß-glucosidase activity by 25% relative

Acid

524
589
527

ns
ns

to the caatinga values.

Soils under algaroba were similar to those of the
natural caatinga. Only LF-N (75%), LF-Po (60%),
available P (120%) and K (105%) were greater than
Mic-P

5.8

6.4
6.3

ns
ns

the caatinga. No differences between caatinga and

algaroba were obtained for the soil microbiological
parameters.
Mic-N

11 ∗∗∗

The paired comparison of the soil chemical val-

28

21
32

comparisons were n=8. Values are the difference between the soils.
7

ues (Table 6) under the tree canopies versus outside

canopies values showed no differences in total nutri-
Mic-C

ent pools. However, microbial biomass C and N, the

19 ∗
156

106
125

ns

more labile pools, were raised by 120% and 150%,

respectively, under the canopies (Table 7). Similar res-
ults were obtained for alkaline phosphatase (120%)
LF-Po

23.7

26.3
38.6

8.5
ns

and ß-glucosidase (140%).

Soil chemical parameters under algaroba were not
different from those outside the canopies 13 years after
LF-N

0.37

0.40

0.18
0.65

planting. However, Prosopis juliflora has been repor-

ns

ted to restore degraded sodic soils in India, to increase

total SOM and available nutrients (Bhojvaid and Tim-
LSDb
Test

ta

ns = non significant.

mer, 1998) and to increase N levels (Tiedemann and

Klemmedson, 1973). Prosopis juliflora has the po-
Caatinga (C4)
Algaroba (A)
Land cover /
Comparison

Buffel (BA )

C4 / A / BA

tential to establish a symbiosis with N-fixing rhizobia

A / BA

b =0.05.

(Diagne and Baker, 1994). Nitrogen fixation by trees

results in an increase of litter N and of N-cycling and
accumulation under the canopies (Young, 1997). It
also produces lower 15 N-abundance as compared with
non-fixing species exploiting the same soil (Nyberg
and Högberg, 1995). However, we found no signific-
ant differences in N balances or 15 N abundances (data
68
not shown). Diagne (1992) reported that rhizobium
strains compatible with Prosopis juliflora did not ex-
ist in soils of southern Senegal; perhaps conditions in
semi-arid NE Brazil are similar. It is also likely that N
fixation is impaired by the low moisture availability in
NE Brazil.
Conclusions
The native trees traditionally preserved in agricultural
landscapes of semiarid NE Brazil, were associated
with islands of fertility in planted pasture, as already
reported by Menezes et. al. (1999). The levels of
fertility and biochemical indicators of soil quality
were particularly high under the evergreen joazeiro,
in comparison with both the introduced pasture and
the original caatinga. This raises questions about the
mechanisms of establishment and maintenance of the
islands of fertility. The observed high fertility levels
under isolated joazeiros are best explained by a soil–
tree interaction in which the low landscape position
with its higher moisture availability facilitates nutrient
recycling by the evergreen joazeiro. The elimination
of competition by forest clearing may also enhance the
effect of isolated trees. The high nutrient and organic
matter contents and biological activity levels under
these trees indicate that the tradition of preserving nat-
ive species in pasture and farm lands may reflect a
selection for species with high capacity for soil im-
provement. Particularly the integration of joazeiro,
which does not have the added value as a produ-
cer of marketable fruit, into silvo-pastoral systems
may be related to its beneficial effects on soil quality
and sustained productivity. Therefore, the search for
indigenous alternatives to the main agroforestry spe-
cies may provide viable alternatives for improved land
management and the conservation of biodiversity. The
observed increase in total P under joazeiro was sub-
stantiated by comparisons with a number of vegetation
covers in spatial relationships. From this, it appears
possible that the mature tree in an alluvial landscape
position may have contributed up to 100 mg P kg−1
of surface soil through root capture and recycling of
nutrients.
When native caatinga was entirely replaced with
buffel grass pasture or a buffel - algaroba silvopas-
toral system, only microbial biomass C and N, and
alkaline phosphatase and ß-glucosidase activity de-
clined, while most chemical soil quality indicators
were unaffected. The biological activity under the
trees of an algaroba-buffel grass association was im-
proved over that of buffel grass alone. This may reflect
greater nutrient cycling due to the mix of litter inputs,
since the canopy of the algarobas was so transparent
that it did not diminish grass growth. Algaroba was
not as effective in providing high biological activities
as the preserved native trees, and it was not effect-
ive in raising nutrient or organic matter levels. This
may be due to the young age of the trees relative to
the mature caatinga remnants, or may reflect the weak
performance of algaroba under semiarid conditions.
Microbiological and biochemical parameters re-
sponded more readily to changes in vegetation than
chemical fertility indices. They are important biolo-
gical indices of the impact of vegetation changes and
its associated land quality changes in silvo-pastoral
systems.
Acknowledgements
This study was financed by the InterAmerican Institute
for Global Change Research (IAI) and by the Ca-
nadian International Development Agency. The help
from Prof I.H. Salcedo in the site selection, and
the hospitality and access to field sites granted by
Agropecuaria Jaçana are gratefully acknowledged. We
thank Jackie O. Moir and Taina M. Redl for their
assistance with analyses.
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