Applied Soil Ecology 12 (1999) 159±167

The effects of overgrazing on soil microbial community

and fertility in the Chaco dry savannas of Argentina
A. Abrila,*, E.H. Bucherb
a
Departamento de Recursos Naturales, Laboratorio de MicrobiologõÂa AgrõÂcola, Facultad de Ciencias Agropecuarias,
Universidad Nacional de CoÂrdoba, Casilla de Correos 509, 5000, CoÂrdoba, Argentina
b
Centro de ZoologõÂa Aplicada, Universidad Nacional de CoÂrdoba, Casilla de Correos 122, 5000, CoÂrdoba, Argentina
Received 13 April 1998; accepted 3 November 1998

Abstract

We identi®ed and measured changes in soil characteristics, nutrient availability and microbial activity to a gradient of grazing
intensity in the Chaco region of northwestern Argentina (248 430 S and 638 170 W). Three sites were selected for comparison:
1. highly restored (no grazing for 20 years);
2. moderately restored (eight years of restoration); and
3. highly degraded (a typical Chaco campesino dwelling, extremely overgrazed).
On each site two sets of soil and litter samples were taken during the dry and wet season, respectively. The following
parameters decreased as grazing intensity increased: soil moisture (from 4.5% to 2.25%), organic matter (from 4.68% to
1.45%), and nitrogen content (from 0.28% to 0.14%). Salt content increased from 1.8 to 5.0 d smÿ1. Microbial activity ranged
from 0.89 mg CO2 gÿ1 7 dayÿ1 at the restored sites to 0.22 mg CO2 gÿ1 7 dayÿ1 at the highly degraded site. The seasonal
variations in density and activity of microorganisms increased from the highly restored to the highly degraded site, probably as
a response to an increased lack of humidity. The cellulolytic and nitri®er groups were the most affected, whereas the
ammoni®er and free-living nitrogen-®xing organisms decreased in the highly degraded site only. Nitrogen ®xation was more
intense at the moderately restored site followed by the highly degraded site. The observed values are interpreted as resulting
from the interaction between organic matter availability (as energy source) and nitrogen de®ciency. Our results suggest a
strong in¯uence of overgrazing on the soil fertility, as well as on the soil ability to buffer water stress during the dry season.
When compared with other savannas of the world, soils at the restored sites show high soil nutrient and organic matter levels,
probably as a result of high phosphorous availability resulting from their recent geological origin. # 1999 Elsevier Science
B.V. All rights reserved.

Keywords: Savannas; Chaco; Overgrazing; Nutrients; Soil microorganisms; Argentina

1. Introduction

Ecosystem degradation due to overgrazing in

*Corresponding author. Tel.: +54-51-334116; fax: +54-51- semi-arid savannas is a very pervasive and well-
544118; e-mail: aabril@agro.uncor.edu known process over the world, being responsible for

0929-1393/99/$ ± see front matter # 1999 Elsevier Science B.V. All rights reserved.
PII: S 0 9 2 9 - 1 3 9 3 ( 9 8 ) 0 0 1 6 2 - 0
160 A. Abril, E.H. Bucher / Applied Soil Ecology 12 (1999) 159±167
increasing poverty and emigration of the rural popula-
tion, particularly in underdeveloped countries
(Skarpe, 1992). A dramatic example of that can be
seen in the Chaco region of South America, a vast
semi-arid plain of more than 1.2 million of km2
extending over parts of Bolivia, Brazil, Paraguay
and Argentina (Bucher, 1982).
In large portions of the Chaco, the original vegeta-
tion dominated by a mosaic of forest, woodlands,
savannas and grasslands has been drastically altered
after a short, but intense period of overgrazing that
followed introduction of cattle by the Europeans.
Grasslands disappeared rapidly, being replaced by a
dense, thorny shrubland dominated by several species
of Acacia and Mimosa (Morello and Saravia Toledo,
1959; Bucher and Scho®eld, 1981). This process of
shrub encroachment is among the most intense
recorded in savannas (Huntley and Walker, 1982).
At present, it still continues expanding into the rela-
tively untouched Chaco regions of Bolivia and
Paraguay.
Although the overall impact of overgrazing in the
Chaco is well documented, much less is known about
the changes induced by overgrazing at the soil level.
However, an adequate understanding of the connec-
tion between overgrazing and both the nutrient cycling
and soil microbial activity is essential to understand
the dynamics of the Chaco savannas. This question is
still more appealing if we consider that the Chaco soils
are in general richer than soils of other subtropical
savannas of the world due to their recent origin
from eolic and ¯uvial materials (Bucher, 1982;
Huntley and Walker, 1982). Comparison of the
effects of overgrazing between nutrient-rich savannas
like the Chaco with nutrient-poor savannas like the
Cerrado in South America or the Miombo in Africa is,
therefore, of particular interest in order to understand
the role played by availability of nutrients in shaping
vegetation and ecosystem processes in savannas
(Huntley and Walker, 1982; Scholes and Walker,
1993).
The objectives of this study are:
1. to identify and measure the changes in soil
characteristics, nutrient availability and microbial
activity resulting from different grazing pressures
in the drier, western portion of the Chaco in
Northwestern Argentina;
2. to compare our results with similar observations
made in other savannas of Africa and South Amer-
ica; and
3. to discuss their implications in terms of range
management and vegetation restoration.
2. Materials and methods
2.1. Study area
The study was conducted at the Los Colorados
Biological Station, located in the province of Salta,
Argentina, 110 km northwest of the city of Joaquin V.
Gonzalez (248 430 S and 638 170 W). The station,
10 000 ha in size, belongs to the FundacioÂn para el
Desarrollo del Chaco (FUDECHA) based at Salta,
Argentina. It is managed by a cooperative research and
development program that integrates researchers from
FUDECHA, the Centro de ZoologõÂa Aplicada (Uni-
versity of CoÂrdoba) and the FundacioÂn Miguel Lillo
(University of TucumaÂn).
Annual rainfall in the area averages 550 mm, con-
centrated during the summer (October±March). The
dry season extends during winter (April±September).
During this period, the water balance is negative,
resulting in a soil moisture de®cit. Mean temperature
of the warmest month (January) and coldest month
(July) are 28.88C and 16.68C, respectively. The exist-
ing literature (Vargas Gil, 1990) describes the local
soils as deep, loam in texture, from eolic and alluvial
origin, neither saline nor sodic. Slope is very slight, of
around 0.5%. Organic matter content is around 1.5%.
Taxonomically they belong to the ``Aridic Haplustoll''
group (USDA Soil Taxonomy), with the following
horizon sequence: A1, B2, B3, C; the B2 horizon
being light structured.
Vegetation at the station is the typical xerophyllous
subtropical woodland of the Western Chaco with a
ground cover of cacti and grasses. Canopy species
include Schinopsis quebracho-colorado (quebracho
colorado) and Aspidosperma quebracho-blanco
(quebracho blanco). The dense 3±4 m high shrub
layer is dominated by species of Acacia, Mimosa,
Prosopis, and Celtis, with grasses intermingled
(Bucher, 1982).
The general area where the ®eld station is located
was heavily grazed, since approximately 1930 (when
 A. Abril, E.H. Bucher / Applied Soil Ecology 12 (1999) 159±167 161

the railroad reached the area) until about 1976, when a

restoration project started. The ®eld station was fenced
and all grazing from domestic herbivores gradually
excluded, allowing the natural vegetation to recover.
The surrounding areas outside the station remained
exposed to continued and severe overgrazing, mostly
by cattle and goats. They are occupied by Chaco
campesinos (``puesteros'') that live in poor mud
houses around an artesian well, the only source of
water during the dry season (Bucher and Scho®eld,
1981). Cattle and goats are free ranging and grazing,
and is not managed, leading to concentric rings of
increasing overgrazing and vegetation degradation
towards the water source (Morello and Saravia Toledo,
1959). Therefore, availability of both recovered and
degraded areas closely located provides a unique
opportunity for long-term monitoring and compara-
tive analysis.

2.2. Sampling design

Three sites were selected for sampling along a 6 km

long gradient of increasing grazing pressure and
environmental degradation (Fig. 1). Otherwise, the
area was homogeneous in terms of soil and vegetation
characteristics.
Site 1: Highly restored (HR). Located at the center
of Los Colorados Biological Station. This site had no
livestock or forest falling in the previous twenty years.
Before, large quebracho trees were cut for wood. At
present, species composition and structure is close to
the original, unexploited forest (Saravia Toledo,
1987).
Site 2: Moderately restored (MR). Located along
the road that connects sites 1 and 3, 2 km from site 1,
and 3 km from site 2. Still within the Los Colorados
reserve, but with a shorter history of restoration, its
vegetation condition being intermediate between the
other two sites. The site was very degraded by over-
grazing and deforestation at the moment of being
fenced eight years ago. There is a noticeable increase
in grass cover and a large number of young trees,
including grazing-sensitive species as Schinopsis
quebracho-colorado.
Site 3: Highly degraded (HD). A typical Chaco
campesino dwelling, known locally as ``puesto'',
located 5 km SE from site 1 (248 430 5000 S and 638
170 2100 W). This site has been under heavy grazing
Fig. 1. Location of the study area and sites in the Los Colorados
Field Station and the adjacent Campo Grande puesto in Salta
Province, Argentina (HR ± highly restored, MR ± moderately
restored and HD ± highly degraded). Dots are smapling locations.
The extent of the Chaco region in South America is shown.
pressure and forest cutting (for timber and charcoal
production) for the past 60 years. Originally cattle
were the dominant herbivore, but were replaced later
by goats, when carrying capacity decreased because of
vegetation degradation. Domestic herbivores have free
access to the only watering point available. Zones of
increasingly degraded vegetation develop around the
watering point, the central area being almost denuded
of grasses and herbs, with only a few trees remaining
(see a detailed description of the ``puesto'' vegetation
in Morello and Saravia Toledo, 1959).
On each site, two sets of samples were taken: the
®rst during the dry season (August 1993) and the
second during the wet season (April 1994). Each set
consisted of thirty soil and litter samples selected at
random along a 200 m linear transect. Each litter
sample was collected form a core of 25 cm2 in area.
Soil samples consisted of soil from 0±20 cm in depth,
of around 300 g each.
162 A. Abril, E.H. Bucher / Applied Soil Ecology 12 (1999) 159±167
2.3. Soil and litter measurements
Soil samples were air dried for 24 h, sieved through a
2 mm mesh, and stored at 48C until processing. For each
soil sample, the following physical and chemical char-
acteristics were measured: soil moisture by dry weight at
1058C, bulk density using the cylinder of Kopeki (Blake,
1982), pH, organic matter by the wet method of Walkley
and Black (Nelson and Sommer, 1982), total nitrogen by
the micro-Kjeldahl technique modi®ed to colorimeter
and spectrophotometer by Apostolatos (1984), N±NO3
with potentiometer (Keeney and Nelson, 1982), extrac-
table phosphorus by the Bray method (Olsen and
Sommer, 1982), salinity by electrical conductivity
(Rhoades, 1982), and soil texture (Day, 1982).
In litter samples we measured: biomass (material
dried at 608C), total nitrogen by the micro-Kjeldahl
technique (Apostolatos, 1984), organic carbon by the
wet method of Walkey and Black modi®ed for plant
material (Nelson and Sommer, 1982), and litter com-
ponents by direct identi®cation of whole leaves.
Soil microbial communities were estimated by
counting cellulolytic, ammoni®ers and nitri®ers, using
the most probable number methods (NMP) in liquid
medium (Alexander, 1982), and free-living dinitro-
gen-®xing microorganisms by counting in Petri dishes
(DoÈbereiner, 1980). Soil respiration was estimated
through CO2 evolution at 288C in samples incubated
for seven days (Alef, 1995). Measurements were made
in the lab under standardized soil moisture conditions
at 60% of ®eld capacity. Nitrogenase activity in soil
was measured using a modi®ed version of the acet-
ylene reduction assay (ARA) (Limmer and Drake,
1996). To estimate the amount of N2 ®xed annually,
we used a conversion factor of 1/3 N2 reduced per
acetylene reduced (Burris, 1974).
3. Results
3.1. Soil characteristics
Soils of the three sites were similar in terms of their
general physical characteristics. Soil texture was simi-
lar, the sand component being slightly higher in the
highly restored site, probably due to the in¯uence of
an old riverbed close to the sampling site (which did
not affect its topography). Extractable phosphorous
and pH were also similar (Table 1).
Therewere speci®c differencesamongsites, however,
most of them resulting from overgrazing and trampling
(Tables 1 and 2). Soil moisture and organic matter
content showed a substantial decrease along the grazing
intensity gradient, particularly, in the highly degraded
site, whereas salinity increased. In accordance with the
observedfall inorganicmatter,nitrogenlevelsdecreased
whereas soil bulk-density increased.
Litter biomass was slightly higher at the restored
site than at the moderately restored site, but non-
existent at the highly degraded site. Litter under trees
was dominated by leaves from woody species (parti-
cularly Schinopsis quebracho-colorado, Aspido-
sperma quebracho-blanco, Celtis sp. and Capparis
spp.), whereas grass and herb materials were predo-
minant in open spaces. Lichens and algae of the genus
Phormidium (Cyanobacteria) were present in the mod-
erately restored site only, suggesting an early step in
the soil recovery process.
3.2. Site and seasonal variations of microbial
populations
Microbial density and activity (measured as soil
respiration) differed between sites, being in most cases
higher at the restored site (Table 2). These differences
were smaller during the wet season. The seasonal
variations in microbial density and activity between
the wet and dry season were small and non-signi®cant
at the highly restored site, but much more important at
the moderately restored site and particularly at the
highly degraded site, probably as a response to an
increased loss of humidity in the unshaded, bare soils
during the dry season.
The observed decrease in microbial density during
the dry season varied in intensity among functional
groups. The cellulolytic and nitri®er groups were the
most affected (probably due to their lack of resistance
forms and reduced diversity), whereas a reduction in
the ammoni®er and free-living nitrogen-®xing groups
was observed in the highly degraded site only, under
extremely adverse conditions (Table 2).
3.3. Soil nitrogen fixation
The number of free-living nitrogen ®xing organisms
was similar in the highly and moderately restored sites
(8.55  0.43 SD and 7.45  1.11 SD log 10 gÿ1,
 A. Abril, E.H. Bucher / Applied Soil Ecology 12 (1999) 159±167 163

Table 1
Soil and litter characteristics at the three study sites (average of all seasonal samples)

Highly restored (site 1) Moderately restored (site 2) Highly degraded (site 3)

Mean SD Mean SD Mean SD

Soil moisture (%) 4.5 1.30 a 2.05 0.3 b 2.25 1.00 b

Bulk density (g cmÿ3) 0.91 0.11 c 1.09 0.11 b 1.22 0.13 a
pH 6.97 0.32 a 6.94 0.17 a 6.95 0.16 a
Organic-matter (%) 4.68 0.08 a 3.05 0.34 b 1.45 0.10 c
Total nitrogen (%) 0.28 0.03 a 0.18 0.13 b 0.14 0.08 b
N±NO3 (ppm) 14.6 1.30 b 24.0 1.51 a 8.50 1.33 c
Extractable P (mg kgÿ1 ) 52.5 9.52 a 45.5 8.01 a 51.8 7.21 a
Salinity (ds mÿ1) 1.8 1.8 1.1 1.1 5.0 5.0
Texture Sandy loam Coarse silt-loam Silty-clay loam
Litter biomass (kg haÿ1) 4.779 0.31 a 3.869 0.40 a 0.00 0.0 b
Litter total nitrogen (%) 2.03 0.80 a 1.68 0.35 a
Litter C/N 20/1 3.9 a 32/1 1.1 a ± ±
Differences between the values of same variable followed by the same letter are not significant at P<0.05.

respectively) with smaller values in the highly
degraded site (5.15  1.01 SD log 10 gÿ1). When
compared in terms of relative abundance with the
ammoni®ers group, the free-living ®xing group was
more abundant in the moderately restored and highly
degraded sites (by a ratio free-living ®xers/ammoni-
®ers of 1.25 and 1.12, respectively), being smaller at
the highly restored site (0.96).
Differences in nitrogen ®xation among sites did not
show a direct relationship with nitrogen de®cit
(expressed in terms of the differences in N content
with the highly-restored site), as could have been
expected. Nitrogen ®xation was more intense at the
moderately restored site, followed by the highly
degraded site, with the lowest values at the highly
restored site (Fig. 2). We found no correlation
between the amount of nitrogen ®xed and the number
of free-living nitrogen-®xing organisms at each site
(p<0.05 level), suggesting that nitrogen-®xing activity
by free-living microorganisms may increase when
nitrogen is scarce. There was no indication that nitrate
concentrations could reach high enough levels to
repress nitrogenase activity in any of the study sites,
given that there was no correlation between soil nitrate
content and nitrogen ®xed.
4. Discussion
4.1. The impact of overgrazing in soil dynamics
The striking differences found between sites in
terms of soil humidity, salinity, organic matter, nitro-
gen content and microbial activity clearly indicate that

Table 2
Seasonal differences in the soil microbial community at the three study sites

Highly restored (site 1) Moderately restored (site 2) Highly degraded (site 3)

Wet season Dry season Wet season Dry season Wet season Dry season
ÿ1
Ammonifiers (log10 g ) 8.6 A 9.1 a 6.7 B 5.3 b 7.6 B $ 1.8 c
Nitrifiers (log10 gÿ1) 3.8 A 2.3 a 3.6 A $ 0.1 b 2.8 B $ 0.1 b
Cellulolytics (log10 gÿ1) 7.2 A 5.4 a 7.2 A $ 0.8 b 6.2 B $ 1.4 b
N2 fixing (log10 gÿ1) 8.2 A 8.9 a 8.4 A 6.5 b 7.9 B $ 2.4 c
Soil respiration (mg CO2 gÿ1 7 day)ÿ1 0.85 A 0.89 a 0.58 B 0.66 b 0.40 C $ 0.22 c
$: significantly different at P<0.05. Differences between values of same variable followed by the same letter (capital letter: wet season; small
letter: dry season) are not significant at P<0.05.
164 A. Abril, E.H. Bucher / Applied Soil Ecology 12 (1999) 159±167

Fig. 2. Relationship between N deficit, organic matter availability and N2 fixed (kg haÿ1 yÿ1) according to site conditions in the studied
Chaco soil. N deficit is expressed as the difference in percentage between the value found at the highly restored site (which is assumed to
correspond to the no deficit situation) and the other sites; organic matter availability is expressed as a percentage of the highly restored sites
value (assumed to be 100%). Notice that N2 fixation is maximum in the combination of nitrogen deficit (that stimulates N2-fixing activity) and
adequate availability of organic matter as energy source for highly demanding metabolic process as N2 fixation.

overgrazing in the Chaco has a profound impact on
soil fertility. Overgrazing leads to soil compaction,
decreased water retention, increased salinity, and loss
of some nutrients, particularly nitrogen.
Our results also suggest a strong in¯uence of over-
grazing and its associated changes in vegetation con-
dition on the soil ability to buffer water stress during
the dry season. Of particular interest is the fact that in
the restored area, microbial populations and soil
respiration remain at about the same level along the
wet and dry season, probably as a response to the
buffering effects of a high organic matter content (and
the associated water-retention capability) combined
with more shaded environment due to a greater
plant cover. In contrast, microbial activity is
highly seasonal in degraded areas. From the point
of view of soil dynamics, conditions at the restored
site are much closer to those found in a non-seasonal
forest than in a typical savanna, the opposite being the
case at the degraded site. Our ®ndings contrast with
the widespread idea that biological processes are
almost exclusively seasonal in semi-arid regions
(Bezuidenhout, 1978; Mazzarino et al., 1991), being
in agreement with previous work on the in¯uence of
vegetation cover on the biological activity in soils of
the western Chaco of Cordoba, Argentina (Abril et al.,
1993).
Restoration of the original soil conditions after
removal of cattle seems to be a slow process in the
Chaco, as suggested by the fact that conditions in the
moderately restored site after eight years of enclosure
are closer to the overgrazed area than to the fully
restored site. Litter consumption by cattle associated
with intense overgrazing (Morello and Saravia Toledo,
1959) may be one of the key factors limiting soil
recuperation under savanna conditions (Newbould,
1989). It may also affect nutrient cycling by depriving
soils of nutrients captured by the deep root system of
Chaco trees, or capture from precipitation inputs (such
as an increased cation-exchange capacity in the soil
beneath trees, and an increased soil±moisture±reten-
tion capacity at these sites) resulting in preferential
enrichment around trees (Kellman, 1979; Abril et al.,
1993).
 A. Abril, E.H. Bucher / Applied Soil Ecology 12 (1999) 159±167
4.2. Factors affecting nitrogen fixation by free-living
organisms
We expected that nitrogen ®xation by free-living
organisms should be highest at the highly degraded
site, considering that nitrogen ®xation usually
becomes particularly apparent in successional com-
munities (in this case, the early stage along the
restoration gradient) (Silver et al., 1996). However,
the fact that the highest level of nitrogen ®xed was
found in the transition site might suggest that N2
®xation is adversely affected at the most degraded
areas.
The observed values could be interpreted as
resulting from the interaction between nitrogen
de®ciency and organic matter availability, considering
that other factors as pH or nitrate content were
not limiting in our case (Nohrstedt, 1982; Limmer
and Drake, 1996). When nitrogen and organic
matter availability are high, as in the restored
site, nitrogen ®xation from the atmosphere should
decrease as the microorganisms shift to alternative,
energetically less-expensive sources of N in the
soil. On the other extreme, when nitrogen availability
and organic matter are low (as in the highly degraded
site), nitrogen ®xation would also be limited, despite
its scarcity, by the lack of carbon-related energy
sources (organic matter) required for a highly
energy-demanding process such as N2 ®xation
(Fig. 2).
We conclude, therefore, that nitrogen ®xation by
free-living microorganisms is an important process in
the Chaco, particularly in degraded areas. However,
lack of organic matter in the soil resulting from
overgrazing and litter remotion in severely degraded
areas may affect N ®xation and result in a substantial
delay in the soil recovery process. As long as
energy from organic matter becomes again available
(after cattle is removed and vegetation recovers), the
increasing activity of N2 ®xing microorganisms
allows other microorganisms to multiply, the
resulting biomass becoming the most important nitro-
gen source in the Chaco soils, as mentioned by
Mazzarino et al. (1991, 1997). Further research would
be needed to increase our understanding of these key
aspects of the Chaco nutrient dynamics, with impor-
tant implications in terms of ranching and restoration
practices.
165
4.3. The Chaco as a nutrient-rich savanna
Our results indicate that soils under the restored or
original Chaco forest are very fertile, in agreement
with reports from several sites within the same sample
region (Casas et al., 1978). Contrasting with our
®ndings, there is a widespread belief that Chaco soils
are limited in terms of their fertility (Vargas Gil,
1990). We presume that this underestimation results
from assuming that samples obtained from degraded,
overgrazed sites (which are predominant in the Chaco)
are representative of average soil conditions from the
original forest or restored areas.
The Western Chaco soil nutrient contents are rela-
tively high when compared with other savannas of the
world, most of which are located in soils of much older
origin, having being exposed to long periods of weath-
ering and leaching (Huntley and Walker, 1982). The
values found in the Chaco were of similar order, but
somewhat higher than those recorded at the Nysvley
Savanna in South Africa (organic matter: 4.1%; nitro-
gen: 0.18% and phosphorus: 32 ppm) (Scholes and
Walker, 1993). They were also higher than the typical
soils of the Brazilian Cerrado (organic matter: 1.3%;
nitrogen: 0.06%, phosphorus: 10 ppm) (Eiten, 1982).
The Cerrado is a typical example of a wet, nutrient
poor savanna, where soils are of much older origin.
The same applies to the Australian Savannas (organic
matter: 1.7%; nitrogen: 0.08%, phosphorus: 10 ppm)
(Walker and Gillison, 1982; McKeon et al., 1991).
It is very likely that high nutrient availability (par-
ticularly phosphorous) should play a key role in
allowing nitrogen ®xation (by both free-living and
symbiotic organisms) and organic matter production
in the Chaco, which in turn shapes vegetation structure
and function, including response to overgrazing. For
example, the Western Chaco is dominated by dense
shrubs with thorns and small compound leaves and
high palatability grasses, characteristics that clearly ®t
with the pro®le given by Huntley (1982) and Scholes
and Walker (1993) for nutrient-rich savannas. It is also
possible that the comparatively very intense shrub
encroachment process observed in the Chaco as
response to overgrazing may also be linked to soil
richness, especially if we consider that most of the
invading species are leguminous species with nitro-
gen-®xing capability and (probably) high require-
ments of phosphorous and other nutrients.
166 A. Abril, E.H. Bucher / Applied Soil Ecology 12 (1999) 159±167
Acknowledgements
The authors are grateful to the staff at the Micro-
biology Department, Faculty of Agronomical
Sciences, University of CoÂrdoba for technical advice.
We are also grateful to Dr. Lilian Frioni for critical
reading of the paper. Financial support was obtained
from the Robert D. and Katherine T. Mac Arthur
Foundation, the W. Alton Jones Foundation, and the
Consejo de Investigaciones Cientõ®cas y TecnoloÂgicas
de la Provincia de CoÂrdoba (CONICOR).
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