Catena 75 (2008) 288–296

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Grazing-induced spatial variability of soil bulk density and content of moisture,

organic carbon and calcium carbonate in a semi-arid rangeland
Ilan Stavi a,1, Eugene David Ungar b,2, Hanoch Lavee a,3, Pariente Sarah a,⁎
a
Laboratory of Soil and Geomorphology, Department of Geography and Environment, Bar Ilan University, Ramat Gan 52900, Israel
b
Department of Agronomy and Natural Resources, Institute of Plant Sciences, Agricultural Research Organization – the Volcani Center, Bet Dagan 50250, Israel

a r t i c l e i n f o a b s t r a c t

Article history: The landscape of many semiarid rangelands is characterized by a two-phase, shrub–intershrub vegetation
Received 10 February 2008 mosaic, each phase having different soil properties. However, this broad subdivision groups together types of
Received in revised form 14 July 2008 intershrub surface cover that may also differ in their soil properties and play important roles in ecosystem
Accepted 23 July 2008
functioning. In the northern Negev region of Israel, we examined the soil properties associated with flock
trampling routes and rock fragment clusters, as well as those associated with the remainder of the intershrub
Keywords:
area and shrub patches. Moisture content, organic carbon content, bulk density and calcium carbonate
Grazing
Shrub
content of the soil were determined for the above four types of cover, inside and outside long-term grazing
Trampling route exclosures. Soil was sampled in the peak of the growing season and in the end of the dry season, on a north-
Bulk density and a south-facing hillside, and from two depths. The shrub patches exhibited the highest soil moisture and
Organic carbon organic carbon contents, and the lowest bulk density and calcium carbonate contents. The trampling routes
Soil moisture showed opposite trends. The rock fragment clusters and the remainder of the intershrub area did not
generally differ and had intermediate values of these properties. Grazing did not have a significant effect on
soil properties at the whole-plot scale, but there were highly significant interactions between grazing and
type of cover. Compared with the former trampling routes in the exclosures, the active trampling routes
outside them had higher bulk density and lower moisture and organic carbon contents. The intershrub area
had higher moisture and organic carbon contents under grazing than in the exclosures. Grazing increased the
spatial heterogeneity of the soil properties examined via the creation of a network of trampling routes on the
hillsides. The routes themselves, which constituted over 20% of the landscape cover, had degraded soil
properties but they led to the improvement of the properties of the remainder of the intershrub area via
functionally important source–sink relationships. The study of the soil of regions in which such networks are
apparent should be duly cognizant of this intershrub subdivision in addition to the widely recognized shrub–
intershrub dichotomy.
© 2008 Elsevier B.V. All rights reserved.

1. Introduction The simplest subdivision of the surface of many semi-arid range-

Soil moisture content is the major driving variable of primary
productivity in semi-arid rangelands (Le Houerou et al., 1988). The
spatial heterogeneity of this resource is fundamental to the function-
ing of these ecosystems, and it is intimately linked to the patchiness of
abiotic and biotic components of the landscape (Noy-Meir, 1973). An
understanding of the soil moisture content associated with the
various types of cover within the landscape and of the redistribution
of water among them is important for the understanding of ecosystem
functioning (Tongway and Ludwig, 2003).
⁎ Corresponding author. Tel.: +972 3 5318808; fax: +972 3 5344430.
E-mail addresses: istavi@yahho.com (I. Stavi), eugene@volcani.agri.gov.il
(E.D. Ungar), laveeh@mail.biu.ac.il (H. Lavee), pariens@mail.biu.ac.il (P. Sarah).
1
Tel.: +972 3 5318808; fax: +972 3 5344430.
2
Tel.: +972 3 9683411; fax: +972 3 9669583.
3
Tel.: +972 3 5318724; fax: +972 3 5344430.
lands is into a two-phase mosaic comprising shrub patches and inter-
shrub areas, with the former acting as sinks for water and other water-
conveyed resources, of which the latter act as sources (Bergkamp,
1998; Lavee et al., 1998; Eddy et al., 1999; Galle et al., 1999). The
quantitative relationships between source and sink areas depend on
several factors, including their respective soil properties such as in-
filtration capacity and the propensity to form mechanical crusts (Carmi
and Berliner, 2008). In the semi-arid northern Negev, Katra et al. (2007)
found that soil moisture content after rain events that initiated
overland flow of water had a patchy pattern, being higher in the shrub
patches than in the intershrub areas. The difference between the two
types of cover diminished with passing time after the rain event but,
nevertheless, the shrub patches exhibited higher soil moisture con-
tents than the intershrub areas.
In addition to soil moisture content, we chose to focus on three
other soil properties – bulk density, organic carbon content and
calcium carbonate content – whose spatial distribution is expected to

0341-8162/$ – see front matter © 2008 Elsevier B.V. All rights reserved.
doi:10.1016/j.catena.2008.07.007
 I. Stavi et al. / Catena 75 (2008) 288–296
be associated with the various types of surface cover and to be affected
by the presence of grazing animals. These soil properties are closely
interrelated and are well understood in terms of the dynamics of plant
litter cycling. Litter reduces direct raindrop impact on the soil, pre-
vents formation of mechanical crusts, and enhances infiltration
(Rostango and del Valle, 1988; Dunkerley and Brown, 1995; Bromley
et al., 1997); it also reduces evaporative losses of moisture from the
soil (Garner and Steinberger, 1989; Tian et al., 1993; Boeken and
Orenstein, 2001). The enhanced soil moisture content increases de-
tritivore abundance and activity, which leads to higher rates of litter
decomposition and incorporation of the resulting humus into the soil
(Warren et al., 1986a). This increases the soil organic carbon content,
reduces bulk density (Thurow et al., 1986; Wilcox et al., 1988) and
increases hydraulic conductivity (Balliette et al., 1986; Wood et al.,
1987). In any specific region the spatial distribution of the calcium
carbonate content indicates the distribution of leaching capacity of the
soil (Shaharabani, 1985; Zwikel, 2004), and its tendency to aggregate;
the higher the calcium carbonate content, the less the aggregation
(Zwikel, 2004). Since soil aggregation affects its hydraulic features, a
feedback loop operates whereby the calcium carbonate content is
both affected by and affects the soil leaching capacity.
Livestock affects soil properties via two main mechanisms: tram-
pling compacts the soil and increases bulk density (Van Haveren, 1983;
Warren et al., 1986b; Trimble and Mendel, 1995); and herbivory
reduces plant cover (Weltz and Wood, 1986; Coughenour, 1991). The
presence of the grazing animal may influence also the source-sink
relations through modification of the landscape mosaic. In the
northern Negev region of Israel close observations of the landscape
surface suggest that treatment of the intershrub area as a single unit
may be an over-simplification. Within the intershrub area are apparent
trampling routes, which we suggested may constitute a separate mi-
crohabitat, i.e., they function differently and their soil properties are
distinct from those of the remainder of the intershrub area. Also
apparent are rock fragment clusters, which may be assembled through
the action of hooves on scattered rock fragments, and which might
constitute another microhabitat because of their effect on the hydro-
logical characteristics of the ground surface (Poesen and Ingelmo-
Sanchez, 1992).
In the present study we asked to what extent the four different
types of cover (shrub patches, trampling routes, rock fragment clusters,
and the remainder of the intershrub area) differ in their soil properties
(bulk density, and moisture, organic carbon and calcium carbonate
contents). In addition, by comparing these properties inside and out-
side long-term grazing exclosures, we asked what impact livestock has
upon them. We also asked to what extent the measured soil properties
are interrelated. We hypothesized that: (i) the four types of surface
cover studied differ in their soil properties; (ii) these differences are
modulated by hillside aspect; (iii) grazing increases bulk density and
calcium carbonate content, and decreases soil moisture and organic
carbon contents; and (iv) grazing has the greatest impact on soil pro-
perties of the type of surface cover that experiences the most intense
hoof action (trampling route), and the smallest impact on that which
experiences the least (shrub patch).
2. Materials and methods
2.1. Area description
The research was conducted at the Lehavim Bedouin Demonstra-
tion Farm, located in the northern Negev region of Israel (31°20' N,
34°46' E). This is a hilly, semi-arid area, 350–500 m above sea level,
with mean annual precipitation of about 300 mm that falls between
October and May (Perevolotsky and Landau, 1988; Bar'am, 1996).
Average daily temperatures in the hottest and coldest months (July
and January, respectively) are 25 and 11 °C, respectively. Relative
humidity ranges between 51% in May and 68% in January (Bitan and
289
Rubin, 1991). The lithology is chalk of the Eocene. The soil type is
Brown Rendzina. The soil is shallow, generally not deeper than 20 cm
in open spaces between shrubs and 40 cm under shrubs, except in
rock fissures. The color is dark brown (7.5YR4/3) and the texture is
clay-loamy in which the primary particle size distribution is 30% clay,
40% silt and 30% sand. The dominant clay types are calcite and
montmorillonite, and the stone content is about 15–30% (Dan and
Koyumdjisky, 1979; Zwikel et al., 2007).
The study region, like many other semi-arid areas of the Old World,
has been grazed by flocks of sheep and goats for centuries. The 800-ha
farm was established in 1980, and has since been managed under the
auspices of the Ministry of Agriculture and Rural Development; it is
grazed by flocks of sheep and goats, totaling about 800 head of breeding
stock.
2.2. Surface cover survey
A survey was conducted of the surface cover components of the
study site in January 2005. Three plots of 4 × 4.5 m were randomly
selected on each of an opposing north- and south-facing hillside of slope
15° and 13°, respectively, located near the central corral, in an area
frequented by the flocks and subjected to intense grazing pressure. A
metal frame of 2.0× 1.3 m was constructed with metal wires stretched
across it to create a grid of 260 cells of 0.1 × 0.1 m. The frame was
supported by four telescopic legs which enabled it to be leveled
horizontally. The area of each plot (excluding two central access strips of
0.3 m width) was mapped using six placements of the frame. For each
cell the cover component with the largest surface area was recorded.
2.3. Soil sampling
Soil sampling was conducted on the same pair of north- and south-
facing hillsides used for the surface cover survey. A number of
permanent exclosures, each measuring 10 × 10 m, were established on
these hillsides in 1994, to serve as long-term, ungrazed controls. On each
of the hillsides we delineated three sampling areas, situated at the
middle of each slope. Each sampling area comprised one control ex-
closure and an adjacent area; hence there were a total of 12 expe-
rimental plots. The sampling areas were approximately 40 m apart.
There were two soil sampling periods, the first at the peak of the
growing season (March) and the second at the end of the hot, dry
season (September) of 2005 when the soil and herbaceous vegetation
were completely dry. Total precipitation at the research site from the
start of the rainy season until the spring sampling was 380 mm.
Monthly totals were 26, 119 and 57 mm in October, November and
December 2004, respectively, and 86, 54 and 38 mm in January,
February and March 2005, respectively. The spring sampling started
12 days after the latest rainstorm, and both the spring and summer
samplings extended over three consecutive days during which there
was no rainfall. In order to minimize the impact of dew on soil
moisture content, soil sampling was initiated in the late morning.
Soil was sampled from under each of four types of cover which
were chosen on the basis of their dominance in the surface cover
survey (see Results): shrub (represented by Sarcopoterium spinosum,
which had a high cover on both aspects); flock trampling route (or
former trampling route in the ungrazed plots); rock fragment cluster
(defined as a group of at least five rock fragments N5 cm in size, which
touched each other and were either semi-embedded in the soil or
lying on its surface); and the rest of the intershrub area (i.e., excluding
trampling routes and rock fragment clusters). Herbaceous vegetation
patches were sampled for the intershrub area because they con-
stituted the dominant cover type in that area. For each type of cover,
soil was sampled from seven randomly selected points within each of
the 12 experimental plots. Under the shrub patches soil was sampled
from 5 cm to the west of the central stem cluster. At each point soil
samples were collected from a depth of 0–2 cm and, where soil depth
290 I. Stavi et al. / Catena 75 (2008) 288–296

permitted, 5–10 cm. Within each sampling day, the southern aspect
was sampled before the northern one.
Moisture content was determined gravimetrically (Gardner, 1965),
and organic carbon content by the wet combustion method (Head,
1984). Bulk density was determined by the clod method (Blake, 1965),
and calcium carbonate content with a calcimeter (Loeppert and
Suarez, 1996). The latter method was evaluated and found to be quite
satisfactory in measuring calcium carbonate content of various soils
along a climatic gradient in Israel, including the semi-arid northern
Negev desert (Zwikel, 2004). Soil moisture content, organic carbon
content and bulk density were determined in both sampling seasons.
Calcium carbonate content was determined only in the spring, since it
is relatively stable, with very little variation in the short term (Sarah,
1999; Zwikel, 2004).

2.4. Statistical analysis

Statistical analysis of each of the measured variables was conducted Fig. 2. Typical south-facing hillside at the study site showing shrubby patches,
with the GLM (general linear model) procedure of SAS (SAS Institute, intershrub areas and trampling routes.

1990). For each season and soil depth, a split-plots type of analysis of
variance (ANOVA) was performed, with aspect (north- or south-facing
underneath the canopy of both shrub species. The most dominant com-
slope) and sampling area (fenced exclosure plus adjacent grazed area)
ponent of the intershrub area was herbaceous vegetation (primarily
as the main plots, and type of cover (shrub, trampling route, rock
annual grasses and forbs), which had a very similar cover to that of S.
fragment cluster, intershrub area) as the sub-plots. Factors in the
spinosum. The next most dominant component of the intershrub area
model were: Aspect (1 df); Sampling area within Aspect (4 df; error
comprised flock trampling routes of almost bare, compacted soil which
term for Aspect); Grazing (1 df); Aspect × Grazing (1 df); Grazing ×
traversed the hillsides with a predominantly horizontal orientation, and
Sampling area within Aspect (4 df; error term for Grazing and Aspect ×
were visually recognizable because of their mechanically formed crusts
Grazing); Cover type (3 df); Aspect × Cover type (3 df) and Grazing ×
and the very sparse herbaceous cover (Fig. 2). The cover of these was
Cover type (3 df). Statistically significant interactions were subjected to
similar on both hillsides and accounted for an average of 21% of the area.
further ANOVA with the SLICE command of PROC GLM. Separation of
Rock fragment clusters were found on both hillsides, though more on
means for Cover type was by Tukey's HSD at the 0.05 probability level.
the south- than on the north-facing slope, and comprised of surface-
Pearson correlation coefficients were calculated to examine the
lying and partially embedded fragments.
relationships among the soil properties.

3.2. Soil moisture content

3. Results

The two sampling seasons were characterized by very large dif-

3.1. Ground surface cover
ferences in soil moisture content. Overall mean percentage water con-
tent (±SE) for spring (March) and summer (September) were 11.2± 0.28
The proportions of the area occupied by the main types of cover
and 2.8 ± 0.04%, respectively, for the shallower (0–2 cm) sampling depth,
encountered in the survey are shown in Fig. 1. The dominant shrub
and 14.1 ± 0.16 and 3.5 ± 0.05% for the deeper (5–10 cm) sampling depth,
species on both aspects was S. spinosum, which is a moderately palatable
respectively.
cushion-like shrub, 20–60 cm in height, and with width ranging from
In the spring, soil moisture content was greater on the north-facing
30 cm for small individuals to more than 100 cm for older, well-
than on the south-facing aspect, with the difference being greater
developed patches. The shrub Coridothymus capitatus – an unpalatable
and more highly significant for the shallower soil depth (north – 13.6%,
cushion-like shrub, 30–40 cm in height, 30–50 cm in width – occurred
south – 9.3%, P b 0.01) than for the deeper soil depth (north – 15.1%,
on the south-facing aspect only. A litter layer of about 1 cm was found
south – 13.1%, P b 0.05). In the summer, no difference was detectable
between the soil moisture contents on the two aspects, at either soil
depth. Neither Grazing nor the Grazing × Aspect interaction had signi-
ficant effects, in the spring or the summer, or at either sampling depth.

Table 1
The gravimetric water content of the soil (%) associated with four types of cover,
according to season and depth of sampling

Sampling season

Spring Summer

Sampling depth (cm)

Cover type 0–2 5–10 0–2 5–10

Shrub patch 15.7 a 16.1 a 3.4 a 4.3 a
Intershrub area 11.6 b 14.0 b 2.6 bc 3.5 b
Rock fragment cluster 11.8 b 13.6 bc 2.7 b 3.6 b
Trampling route 6.8 c 12.8 c 2.4 c 2.9 c

Fig. 1. Relative cover of the main cover types encountered in the ground surface survey Means within a column followed by different letters differ at the 0.05 probability level
according to aspect. according to Tukey's HSD.
 I. Stavi et al. / Catena 75 (2008) 288–296 291

Cover type had a highly significant (P b 0.001) effect on soil Table 2

moisture content in both seasons and at both soil depths. Soil The organic carbon content of the soil (g kg− 1) associated with four types of cover,
according to season and depth of sampling
moisture content was greatest under the shrub patches and lowest
under the trampling routes, with that under the intershrub areas and Sampling season
rock fragment clusters having intermediate values (Table 1). This Spring Summer
ranking was found on both aspects, except for the deeper soil depth on
Sampling depth (cm)
the south-facing aspect in the spring, where the soil under rock
Cover type 0–2 5–10 0–2 5–10
fragment clusters exhibited the lowest soil moisture content, and this
probably accounts for the significant (P = 0.0171) Aspect × Cover type Shrub patch 31.3 a 19.4 a 28.7 a 19.2 a
Intershrub area 21.3 b 15.8 b 20.4 b 15.8 b
interaction obtained for this analysis.
Rock fragment cluster 20.1 b 14.7 b 21.8 b 16.1 b
The interaction Grazing × Cover type was highly significant Trampling route 14.8 c 12.3 c 16.6 c 13.7 c
(P b 0.001) at both soil depths in the spring. For both the ungrazed
Means within a column followed by different letters differ at the 0.05 probability level
and the grazed areas, the soil moisture content at the shallower soil according to Tukey's HSD.
depth differed significantly between all cover types, except the
intershrub area and rock fragment cluster, but that under each cover
type did not differ significantly between the grazed and ungrazed
areas. However, the means for the shrub, rock fragment cluster and and in the summer, respectively). The factor Grazing had no significant
intershrub soil samples were slightly greater in the grazed than in the effect on soil organic carbon content in either season or at either soil
ungrazed area (with the difference being close to significant for the depth. The effect of the Grazing × Aspect interaction was significant
intershrub samples: 12.5 versus 10.6%, P = 0.0552), and those for the (P b 0.05) for the deeper soil depth in the spring: on the north-facing
trampling route were slightly lower (Fig. 3). For the deeper soil depth slope, soil organic carbon content was 15.3 g kg− 1 in the ungrazed area
in both the grazed and the ungrazed areas, the soil moisture content and 16.4 g kg− 1 in the grazed area, whereas on the south-facing slope
under the shrub patches was significantly (P b 0.001) greater than that it was 16.3 g kg− 1 in the ungrazed area and 14.3 g kg− 1 in the grazed
under the other cover types. In addition, in the grazed area the soil area.
moisture content of the intershrub component was significantly Cover type had a highly significant (P b 0.001) effect on soil organic
(P b 0.001) greater than that under the trampling routes. Comparison carbon content in both seasons and at both depths. Organic carbon
between the grazed and ungrazed areas for the same cover type, content was always highest under the shrub patches and lowest under
shows that the trampling route had a higher moisture content in the the trampling routes. All differences among cover types were signi-
ungrazed than in the grazed area (13.7 versus 11.9%, P = 0.0015). ficant except for that between the intershrub areas and the rock
In the summer, the Grazing × Cover type interaction was highly fragment clusters (Table 2).
significant (P = 0.002) for the shallower soil depth only. In the The effect of the Aspect × Cover type interaction was significant at
ungrazed area the soil moisture content was significantly (P b 0.001) both the shallower and the deeper depths (P = 0.0015 and 0.0093,
greater under the shrub patches than under the other three types of respectively) in the spring. At the shallower soil depth, organic carbon
cover, which did not differ from one another. In the grazed area, all the content was lower on the south- than on the north-facing slope for all
cover types differed significantly (P b 0.01) from each other except the cover types except the shrub patch, for which it was similar on the two
intershrub and the rock fragment cluster. As in the spring, comparison aspects. At the deeper depth, organic carbon content was slightly
between the grazed and ungrazed areas for each cover type revealed lower on the south-facing than on the north-facing slope under the
higher moisture contents under the trampling routes in the ungrazed trampling routes and rock fragment clusters, and higher under the
than in the grazed areas (2.7 versus 2.2%; P b 0.001). In the summer, shrub patches and intershrub areas.
differences in soil moisture content were much smaller than in the The effect of the Grazing × Cover type interaction was highly
spring. significant (P b 0.001) at both soil depths in the spring; and significant
(P = 0.0020) at the shallower soil depth and close to significant for the
3.3. Soil organic carbon content deeper soil depth (P = 0.0539) in the summer. Overall, the ranking of
the various cover types with respect to their soil organic carbon con-
The overall mean (±SE) percentage soil organic carbon contents for tent for the four cover types was the same within each combination of
the shallower and deeper soil depths were 21.7 ± 0.39 (n = 624) and soil depth, season, and Grazing; the highest values were found under
15.7 ± 0.22 g kg− 1 (n = 517), respectively. At the shallower depth these the shrub patches and the lowest under the trampling routes (Fig. 4).
contents were higher on the north-facing than on the south-facing However, the difference between the shrub patches and the trampling
slope (23.6 and 20.0 g kg− 1, respectively in both seasons) but these routes tended to be greater in the grazed than in the ungrazed areas. At
differences were not significant (P = 0.0767 and 0.3409 in the spring the shallower soil depth in the ungrazed areas, only the shrub patches

Fig. 3. Mean gravimetric water content of the soil in the spring according to cover type, Fig. 4. Mean organic carbon content of the soil in the spring according to cover type,
grazing treatment and sampling depth. Bars represent one standard error of the mean. grazing treatment and sampling depth. Bars represent one standard error of the mean.
292 I. Stavi et al. / Catena 75 (2008) 288–296

Table 3 Table 4
The bulk density content of the soil (g cm− 3) associated with four types of cover, The calcium carbonate content of the soil (%) in the spring season associated with four
according to season and depth of sampling types of cover, according to depth of sampling

Sampling season Sampling depth (cm)

Spring Summer Cover type 0–2 5–10

Sampling depth (cm) Shrub patch 20.0 c 19.7 b

Intershrub area 20.2 bc 20.3 b
Cover type 0–2 5–10 0–2 5–10 Rock fragment cluster 21.1 ab 20.9 ab
Shrub patch 1.44 c 1.47 b 1.42 c 1.47 b Trampling route 21.4 a 21.5 a
Intershrub area 1.49 b 1.50 b 1.50 b 1.50 a
Means within a column followed by different letters differ at the 0.05 probability level
Rock fragment cluster 1.52 b 1.50 ab 1.49 b 1.50 ab
according to Tukey's HSD.
Trampling route 1.57 a 1.53 a 1.57 a 1.51 a

Means within a column followed by different letters differ at the 0.05 probability level
according to Tukey's HSD. densities were measured under the trampling routes and shrub pat-
ches, respectively, with those under the intershrub areas and rock
fragment clusters having intermediate values. However, in the spring,
none of the differences among the cover types within the ungrazed
differed significantly from the other cover types, in organic carbon
area were significant, whereas all such differences within the grazed
content. In the grazed area, all cover types differed significantly from
area were highly significant (P b 0.001), except for that between the
one another, except for the intershrub areas and rock fragment
intershrub areas and rock fragment clusters (P = 0.8445). The densities
clusters. This was found in both sampling seasons. At the deeper soil
ranged from 1.422 g cm− 3 under the shrub patches to 1.635 g cm− 3
depths, the only difference that was significant in both the grazed and
under the trampling routes. In the summer, bulk density in the un-
the ungrazed areas, and also in both seasons, was that between the
grazed area was significantly (P b 0.001) lower under the shrub patches
shrub patches and the trampling routes. Grazing had a significant
(1.412 g cm− 3) than under the three other cover types (1.494–1.515 g
effect on the organic carbon content of the soil under a given cover type
cm− 3), among which there were no significant differences. In the
for the shallower soil depth of the intershrub area in the spring only:
grazed areas, all differences were significant (P b 0.05) except that
the mean organic carbon content under this cover type was 23.7 g kg− 1
between the intershrub and rock fragment cluster (P = 0.7265). The
in the grazed areas compared with 18.8 g kg− 1 in the ungrazed areas
bulk densities ranged from 1.419 g cm− 3 under the shrub patches to
(P = 0.0038).
1.621 g cm− 3 under the trampling routes. When the grazed and un-
grazed areas were considered together in each season, the bulk density
3.4. Soil bulk density
under the trampling routes in the grazed area was significantly higher
than that under all other cover types. The bulk density under the shrub
The overall mean (± SE) soil bulk densities for the shallower (0–
patches in the grazed area was significantly lower than that under all
2 cm) and deeper (5–10 cm) soil depths were 1.501 ± 0.00443 (n = 624)
other cover types, except that under the shrub patches in the ungrazed
and 1.497 ± 0.00370 (n = 517) g cm− 3, respectively. The factors Aspect,
area.
Grazing and their interaction had no significant effects on soil bulk
density in either season or at either soil depth. Cover type was found
3.5. Calcium carbonate content
to be highly significant (P b 0.001) for both soil depths and both
seasons. Bulk density was always highest under the trampling routes
In the spring the overall mean (±SE) calcium carbonate contents in
and lowest under the shrub patches (Table 3).
the soil at the shallower (0–2 cm) and deeper (5–10 cm) depths were
The effect of the interaction Aspect × Cover type was significant
15.0 ± 0.29 (n = 144) and 14.6 ± 0.31% (n = 127), respectively, for the
(P = 0.0014) in the summer at the shallower depth. Although the
north-facing slope, and 26.3 ± 0.26 (n = 168) and 26.7 ± 0.35% (n = 144),
ranking of bulk density according to cover type was not affected by
respectively, for the south-facing slope. The differences between the
Aspect, the bulk density on the south-facing aspect was slightly lower
two aspects were significant for both soil depths (P b 0.01). The effects
than that on the north-facing aspect under the trampling routes, but
of Grazing and the Aspect × Grazing interaction were not significant
slightly higher under the other three cover types.
for either of the soil depths.
The effect of the interaction Grazing × Cover type was highly
The calcium carbonate content of the soil was affected by cover type
significant (P b 0.0001) at the shallower depth in both seasons (Fig. 5).
at both soil depths (P b 0.001), being highest under the trampling
Within the ungrazed and the grazed areas, the highest and lowest bulk
routes and lowest under the shrub patches, although not all differences

Fig. 5. Mean bulk density of the soil (0–2 cm) according to cover type, grazing treatment Fig. 6. Mean calcium carbonate content of the soil in the spring according to cover type,
and season. Bars represent one standard error of the mean. aspect and sampling depth. Bars represent one standard error of the mean.
 I. Stavi et al. / Catena 75 (2008) 288–296
among cover types were significant (Table 4). However, the effect of the
interaction Aspect × Cover type was significant for both the shallower
and deeper soil depths (P = 0.0117 and 0.0057, respectively), because of
the large changes in the relative magnitudes of the calcium carbonate
content of the four cover types according to aspect (Fig. 6). For the
shallower soil depth on the north-facing slope, the calcium carbonate
content was lowest for shrub patches (14.43%) and similar among the
other three cover types (15.14–15.29%), but none of the differences
among cover types was significant. In contrast, on the south-facing
slope there was greater differentiation between the cover types, with
the highest calcium carbonate content (27.5%) found under the tram-
pling routes, followed, in descending order, by the rock fragment
clusters, shrub patches and intershrub areas (25.5%). Significant dif-
ferences were found between the trampling routes, on the one hand,
and the shrub patches (P b 0.01) and intershrub areas (P b 0.001), on the
other hand, as well as between the rock fragment clusters and the
intershrub areas (P b 0.01). At the deeper soil depth, the calcium car-
bonate contents under the shrub patches were the lowest on both
aspects, whereas that under the trampling route was intermediate on
the north-facing slope and relatively high on the south-facing slope.
The calcium carbonate content ranged from 14.4 to 15.0% on the north-
facing slope, where no differences among the means were significant.
On the south-facing slope, the calcium carbonate content ranged from
25.0% under the shrub patches to 28.3% under the trampling route, and
the latter differed significantly from those under both the intershrub
areas (P b 0.01) and the shrub patches (P b 0.001).
The effect of the interaction Grazing × Cover type was significant
(P b 0.01) only at the shallower soil depth in the spring (Fig. 7). In the
ungrazed area, the calcium carbonate content was significantly lower
under the shrub patches (19.2%) than under the rock fragment clusters
(20.9%; P b 0.05) or the trampling routes (21.2%; P b 0.001). In the grazed
area, the intershrub areas had the lowest calcium carbonate content
(19.8%), which differed significantly (P b 0.01) from that under the tram-
pling route, which had the highest value (21.6%).
3.6. Interrelationships among soil properties
Examination of the Pearson product-moment correlations among
the soil properties revealed a number of highly significant relationships.
The strongest relationships were positive ones between the organic
carbon and moisture contents of the soil. This relationship was exam-
ined separately for each soil depth and each sampling season. The
resulting four correlations were all highly significant (P b 0.001), with the
correlation coefficient being somewhat higher for the shallower depth
(r = 0.72 and 0.63 in spring and summer, respectively) than for the
deeper depth (r = 0.54 and 0.46 in spring and summer, respectively).
Fig. 7. Mean calcium carbonate content of the soil (0–2 cm) in the spring according to
cover type and grazing treatment. Bars represent one standard error of the mean.
293
The organic carbon content and bulk density of the soil were also
strongly related, with the relationship being negative. Since these two
properties did not change appreciably between the two sampling
seasons, the relationship between them was examined separately only
for each soil depth. At both shallower and deeper depths correlations
were highly significant (P b 0.001) and here too the correlation
coefficient was higher for the shallower (r = −0.57) than for the deeper
depth (r = −0.37).
Significant negative relationships were also obtained between the
soil bulk density and moisture content. The relationships were stronger
at the shallower depth (r = −0.38, P b 0.001 in spring; r = −0.52, P b 0.001
in summer) than at the deeper depth (r = −0.12, P b 0.05 in spring; r =
−0.29, P b 0.001 in summer).
The interrelationships with the soil calcium carbonate content
were examined separately for the north- and south-facing aspects and
for the two soil depths. On the north-facing slope, at both depths, this
soil property was negatively related to soil moisture content (r = −0.22,
P b 0.01 for the shallower depth; r = −0.39, P b 0.001 for the deeper
depth). On the south-facing slope, the calcium carbonate and organic
carbon contents of the soil were negatively related (r = −0.29, P b 0.001
for the shallower depth; r = −0.39, P b 0.001 for the deeper depth).
4. Discussion
4.1. Cover type
With the exception of the rock fragment clusters, the different types
of cover selected in this study to represent the surface and above-ground
spatial variability of the hillsides were found to be associated with
distinct sets of soil properties, as hypothesized. A clear distinction was
apparent between the components of the two-phase vegetation mosaic,
i.e., between the shrub patches and the intershrub areas. Underneath the
shrub canopy the bulk density of the soil was relatively low, a finding we
ascribe primarily to the absence of animal trampling. As expected, the
moisture content of the soil in these patches was found to be relatively
high, and presumably derived from the combined effect of a number of
processes, including the accumulation of water overland flow generated
in the intershrub area and shading by the dense shrub canopy. The
organic carbon content of the soil was relatively high at the shrub
patches, and this is consistent with their high litter cover and soil
moisture content because moisture increases the rates of litter decom-
position and organic carbon incorporation into the soil. The increased
organic carbon content enhances the soil's macroaggregate content
(Oades, 1984) and macroporosity and thereby further reduces its bulk
density (Dunkerley and Brown, 1995). The combined effect of these is to
intensify leaching (Sarah, 1999; Zwikel, 2004), an effect that is reflected
in a relatively low content of calcium carbonate. Compared with the
shrub patches, the intershrub areas were found to exhibit lower con-
tents of soil moisture and organic carbon, and higher calcium carbonate
content and soil bulk density.
Our cover survey showed that flock trampling routes constituted
the third largest element, covering over 20% of the landscape. Fur-
thermore, they differed in their soil properties from the shrub and the
remainder of the intershrub area in ways that suggest that they
function as a separate microhabitat. Trampling routes had low vege-
tation cover and high bulk density, which can be attributed directly to
the impact of intensive animal traffic: hoof action damages and
detaches tissue from growing plants (Pande and Yamamoto, 2006) and
compacts the soil surface (Warren et al., 1986a, b). The reduction in
vegetation cover enhances splash impact, mechanical crust formation
and surface sealing (Wilcox et al., 1988; Weltz et al., 1989; Bari et al.,
1993). Compaction also smoothes the soil surface (Nash et al., 2003,
2004). Low vegetative cover, low surface roughness and high bulk
density lead to low infiltration capacity, and consequently the routes
act as net contributors of water to their downslope areas. Using a
standardized soil penetration test as a proxy for infiltration capacity,
294 I. Stavi et al. / Catena 75 (2008) 288–296
Stavi et al. (2008) obtained a mean penetration depth of approximately
6 mm at trampling routes, compared to 10 mm for intershrub areas and
18 mm under the canopy of S. spinosum. The very low soil moisture
content of the trampling routes drives a causal chain of low herbaceous
production, litter production and soil organic carbon content, and
greater soil bulk density. The combined effect of these processes is
expected to reduce the soil leaching capacity, a reduction that is
expressed in a high calcium carbonate content.
The soil properties associated with rock fragment clusters did not
differ from those of the intershrub areas. This might be attributable to
the content of the clusters: surface-lying and partly embedded rock
fragments. Soil moisture content is expected to be higher underneath
surface-lying rock fragments because of a mulching effect whereby the
rock fragments reduce splash impact, surface sealing and crusting, and
thereby increase infiltration (Poesen and Ingelmo-Sanchez, 1992;
Poesen et al., 1994; Valentin, 1994), as well as a shading effect which
reduces evaporative losses (Poesen and Lavee, 1994). In addition, water
flow generated on the surface of the rock fragment augments the
amount of water entering the soil underneath it (Poesen et al., 1994).
Partially embedded rock fragments function differently; they seal the
soil surface and reduce infiltration capacity (Poesen and Ingelmo-
Sanchez, 1992; Poesen and Lavee, 1994; Valentin, 1994). Thus, the
opposite effects of surface-lying and partially embedded rock frag-
ments on the soil water balance may cancel out when the two types of
fragments occur together in a cluster. Such opposing effects may also
account for the lack of difference that was found between rock frag-
ment clusters and the intershrub areas in terms of the organic carbon
content of the underlying soil: although there is less primary pro-
duction at rock fragment clusters (negative effect), they are expected to
trap floating litter (positive effect).
4.2. Hillside aspect
A strong plot-level effect of hillside aspect was detected for
moisture and calcium carbonate contents. As expected, moisture con-
tent was lower on the south- than on the north-facing hillside
(Shoshany, 2002), however, this difference was apparent in the spring
but not following the hot, dry summer months that preceded the
summer sampling. The calcium carbonate content of the soil was much
lower on the north- than on the south-facing hillside. As both aspects
in the research site receive similar amount of rain (Sarah and Lavee,
unpublished data) this is attributable to the relatively high rate of
calcium carbonate leaching that results from the improved soil
aggregation (Stavi et al., unpublished data) and the relatively low
evaporation on the northern hillside. Although there was no plot-level
effect of aspect on the organic carbon content of the soil, cover type
differences were modulated by aspect, as evidenced by the significant
interaction between aspect and type of cover in the spring. A higher
organic carbon content was detected at the north-facing aspect for all
cover types except the shrub patch.
4.3. Grazing
Our results suggest that generalizations about the effect of grazing
on soil properties at the whole plot level are not justified since no such
effects were detected. The effects of grazing need to be examined at
the finer spatial scale of the individual cover type. At this scale, grazing
did increase bulk density and reduce moisture and organic carbon
contents, as hypothesized, but almost invariably only at the trampling
routes.
On the whole, grazing had the highest impact on the most exposed
type of cover, i.e., the trampling routes, and almost no significant
effects on the properties of the soil under the most mulched cover –
the shrub patch. In the shrub patches, presence or absence of the
grazing animal had no effect on the bulk density of the soil, which is
consistent with our impression from the field that sheep and goats do
not trample the area covered by the dense and thorny canopy of S.
spinosum. This is in contrast to Golodets and Boeken (2006) who,
working at a different site in the northern Negev region, found that
sheep trampled the soil mounds under the shrub Noaea mucronata.
This difference is certainly related to the very different canopy
architectures of S. spinosum and N. mucronata, and well illustrates the
danger of broad generalizations. Although we observed the foliage of
S. spinosum to be browsed by both sheep and goats – which might be
expected to alter the soil water balance via changes in evapotran-
spiration losses – there was no significant effect of grazing on the
moisture content of the soil under the shrub patches, except for a
small reduction at the deeper soil depth in the summer. Browsing
might also be expected to influence the organic carbon content of the
soil under the shrub patch by reducing the amount of litter produced,
but no consistent effect of grazing was detected: there was a reduction
in the organic carbon content at the deeper depth in the spring and an
increase at the shallower depth in the summer.
In the intershrub areas, the soil moisture content was greater in the
grazed than in the ungrazed areas. We attribute this to the trampling
route acting as a greater net contributor of water to its downslope area
in the grazed than in the ungrazed areas. A further contributory factor
to the observed difference in moisture content may be reduction of
transpiration losses of moisture in the grazed area. A similar pheno-
menon was observed in the semi-arid Judean Desert of Israel by Sarah
and Lavee (personal communication) who found that the soil moisture
content of grazed intershrub areas was higher than that of non-grazed
plots during the growing season.
Interestingly, the organic carbon content of the soil in the intershrub
areas was also greater in the grazed than in the ungrazed area. Even
though the greater supply of moisture to the intershrub area in the
grazed area might be expected to increase primary production and litter
production, the visibly large difference between the grazed and
ungrazed plots, in standing herbaceous biomass at the end of the
growing season, indicates strongly that much more litter is expected to
be available in the ungrazed plots. A possible explanation might be
enhanced rates of litter decomposition in the presence of the grazing
animal. Because decomposition rates are positively related to soil
moisture content (Raiesi and Asadi, 2006; Yan et al., 2007), which was
higher in the grazed than in the ungrazed intershrub areas, the litter
decomposition rate and hence the soil organic carbon content were also
increased. This was indicated also by the strong positive interrelation-
ship between soil moisture and the organic carbon contents. The bulk
density of the soil of the intershrub areas did not differ between the
grazed and ungrazed areas, which is not surprising, since a major part of
the trampling load is concentrated along the trampling routes.
Prevention of grazing for over a decade reduced the contrast in the
ground surface features and soil properties between the former
trampling routes and the remainder of the intershrub area. In the
control exclosures, trampling routes were much less recognizable due
to the almost complete disappearance of the mechanical crusts. In
both measurement seasons the soil bulk density at the shallower
depth under the trampling routes was significantly lower inside the
exclosure than outside it, which would be expected to increase the
infiltration capacity and soil moisture content of the inside trampling
routes. The latter was indeed significantly greater in the exclosure
than outside it, for both soil depths and in both measurement seasons.
The calcium carbonate content at the deeper depth under the
trampling routes was lower in the exclosure than outside it. Greater
soil moisture content would be expected to enhance herbaceous
colonization, litter production and decomposition, and, thereby, to
cause an increase in the organic carbon content of the soil. Both our
observations and data confirmed these expectations: herbaceous
cover was larger in the former than in the actual trampling routes and
the organic carbon content of the soil under the trampling routes was
indeed significantly greater in the exclosure than outside it, at both
soil depths and in both measurement seasons.
 I. Stavi et al. / Catena 75 (2008) 288–296
4.4. Implications
The presence of livestock created a network of trampling routes
which accounted for over 20% of the surface area of the landscape.
These routes had soil characteristics which were distinct from the
remainder of the intershrub area and which indicated that they play an
important role in ecosystem functioning by influencing the spatial
redistribution of resources at the patch scale. Such non-trophic effects
are readily subsumed under the rubric of the ecosystem engineer
(Jones et al., 1997; Wilby et al., 2001): an organism that regulates the
productivity of other organisms by controlling their resource avail-
ability or by modifying their habitat (Jones et al., 1994). This pers-
pective on animal presence, and trampling in particular, was
propounded by Golodets and Boeken (2006) who found that grazing
increased the proportion of bare soil and decreased microphytic crust
cover, although they did not report on the presence of flock trampling
routes in their study environment. It remains to be seen how wide-
spread a phenomenon the trampling route is in the semi-arid range-
lands of the world, and with what soil, landscape and management
features it tends to be associated. Where present, it needs to be studied
separately in order to better understanding the eco-geomorphic
impacts of livestock presence on these ecosystems.
5. Conclusions
The distinct types of surface cover of hilly, semi-arid rangelands
investigated in this study tend to be associated with distinct sets of soil
properties. Although our findings are consistent with the paradigm of a
two-phase mosaic comprising shrub patches and intershrub areas, it is
important to recognize trampling routes as a separate microhabitat
within the intershrub area. The soil properties of the shrub patches and
the trampling routes lie at opposite ends of the scale, and those of the
intershrub areas fall between them. The study of the soil of regions in
which such networks are apparent should be duly cognizant of this
intershrub subdivision and the effect it may have on the redistribution
of resources and primary production at the patch scale.
Acknowledgements
We acknowledge the support of The Doctoral Fellowships of Excel-
lence of Bar-Ilan University, The Soil Advisory Board and The
Rangelands Advisory Board of The Ministry of Agriculture and Rural
Development and the KKL (JNF). We are grateful to Helena Zhevelev,
Nathan Fragin, Dani Barkai, Rafi Yonatan, Hagit Bar'am and Ezra Ben-
Moshe for their helpful assistance in the field and laboratory work. We
thank two anonymous reviewers for their helpful comments on the
manuscript.
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