ARTICLE IN PRESS

Flora 200 (2005) 49–64

www.elsevier.de/flora

Vegetation–environment relationships and classification of the seasonal

savannas in Venezuela
Zdravko Baruch
Dpt. Estudios Ambientales, Universidad Simón Bolı´var, Apartado 89000, Caracas 1080, Venezuela

Received 25 February 2004; accepted 6 June 2004

Abstract
The seasonal savannas dominated by the C4 grasses of the genus Trachypogon, are widespread in northern South
America. In Venezuela, they extend from the central lowland Llanos to intermediate elevations in the Coastal and
Andean Mountains and to the Guiana Plateau (Gran Sabana) in the south. This study aims to classify these savannas
and to understand the environmental factors that determine their composition and structure. Thirty-seven sites were
sampled, plant cover and density were measured in 376 one square meter quadrats and the importance value index
(IVI) was calculated. Climate data were obtained from climatological stations and soil properties were analyzed. The
results were classified by clustering and TWINSPAN. Floristic and environmental data were ordered with the
canonical correspondence analysis (CCA).
Trachypogon savannas are heterogeneous, both floristically and environmentally. An altitudinal gradient of

2000 m separates the intermediate elevation Coastal Mountains (Type I) and the Guiana savannas (Type II) from the
lowland Llanos. In the lowlands, sparsely covered and treeless communities on extremely oligotrophic and sandy soils
(Type IV) differ structurally and floristically from woody savannas (Type III). The floristics and structure of the
savannas respond directly or indirectly to elevation and water and nutrient availability. These responses were
evidenced by: (a) floristic composition, as Neotropical lowland savanna species merged with those of the Andean sub-
páramo in Type I savannas or with those from the Guianan flora in Type II savannas. (b) Species richness, which
ranged from an average of 13.8 per site in sandy Type IV savannas to 38.9 per site in Type I intermediate elevation
savannas. (c) Community physiognomy, which changes from an homogeneous herbaceous matrix with dispersed trees
in Type III savannas to a shrubby Type I savannas to ‘‘treeless’’ Types II and IV savannas. (d) Plant cover which
varied from 93.8% in the fertile Type I savannas to around 80% in the sandy oligotrophic savannas of Types II and
IV. Each savanna type is related to one of the major geomorphological regions of Venezuela.
r 2005 Elsevier GmbH. All rights reserved.

Keywords: Canonical correspondence analysis; Classification; Conservation; Diversity; Llanos; Neotropics; Ordination

Introduction

Savanna is one of the world’s largest (ca.

15  106 km2) and most productive biomes (Melillo
Fax: +58 212 9063064. et al., 1993; Saugier et al., 2001). In Venezuela, it
E-mail address: zbaruc@usb.ve (Z. Baruch). extends over 220,800 km2, or 24.5% of the country

0076-9511/$ - see front matter r 2005 Elsevier GmbH. All rights reserved.
doi:10.1016/j.flora.2004.06.001
 ARTICLE IN PRESS
50 Z. Baruch / Flora 200 (2005) 49–64
Fig. 1. Map of Venezuela showing the sampling sites and the
savanna types. Numbers correspond to sites named in Table 1.
Scale bar is approximate.
(Huber, 1987). The major continuous savannas are the
central Orinoco Llanos which are limited by the Andean
and Coastal Mountains to the west and north,
respectively (Fig. 1). This ecosystem penetrates deeply
into Colombia where it is known as Llanos Orientales.
Another large savanna area lies in the Guiana Plateau
(Gran Sabana) in the southeast of the country (Fig. 1)
which grades towards the Rupununi-Rio Branco savan-
nas of Guyana and northeastern Brazil. Smaller savanna
patches are found in the Venezuelan Amazon State and
at intermediate elevations (700–2000 m asl; henceforth
named highlands) in the Coastal Mountains. Physiog-
nomically, the savannas are assembled by a low stature
(1–1.5 m) herbaceous matrix of largely C4 grasses and
sedges with a variable proportion of woody plants.
Functionally, they have relatively low primary produc-
tivity by unit area, attributed to unfertile soils, seasonal
droughts and frequent fires (Frost et al., 1986; Medina
and Silva, 1990). In Venezuela, the commonest savannas
are the seasonal type (sensu Sarmiento, 1983) due to
strongly defined dry and wet seasons. They are
dominated by the C4 tussock grasses Trachypogon
plumosus (H. & B.) Nees or T. vestitus Anders (Ramia,
1967; Sarmiento, 1983).
The structure and composition of several local
Trachypogon savannas in Venezuela have been described
previously, at variable levels of detail and with widely
contrasting methods (Blydenstein, 1962; Huber, 1995;
Ramia, 1967; San José et al., 1998; Sarmiento and
Monasterio, 1969; Susach, 1989; Vareschi, 1970; Velás-
quez, 1965). This study differs from the previous by: (a)
it covers all major regions with Trachypogon savannas,
including those south of the Orinoco and, (b) the
sampling and analysis were based on uniform floristic,
numerical and standardized methods based on measure-
ments of species cover and density. This study is timely
because the increasing human pressures on these
savannas compromise their integrity, diversity and
function. Neotropical savannas are being replaced by
crops and forages to increase their productivity (Klink
et al., 1993; Thomas et al., 1990). Oil is being exploited
in the Venezuelan and Colombian Llanos with the
predictable pollution damage. Also, spontaneous inva-
sion of several African grasses is displacing native
savannas in Venezuela and Brazil (Baruch, 1996;
Filgueiras, 2002; Pivello et al., 1999; Williams and
Baruch, 2000). In the near future, Neotropical savannas
will also be affected by increased atmospheric CO2 and
altered rainfall regime (Hoffmann and Jackson, 2000;
Hoffmann et al., 2002) with unpredictable consequences
on savanna composition, physiognomy and stability.
To evaluate the degree of savanna disturbance or
replacement and to offer a base line for restoration
attempts, an understanding of the factors that determine
the savanna’s composition and structure is needed. The
objectives of this study were: (a) To relate the floristic
composition and structure of the savannas with the local
climates, soils and disturbances, and (b) to obtain an
ecologically meaningful classification of the Trachypo-
gon savannas in Venezuela.
Study area
Trachypogon savannas extend over heterogeneous
areas with large variations in topography, climate,
geology, soils, and disturbance regimes (Fig. 1). The
elevation of the study sites ranges between 0 and

2000 m asl (Table 1). Air temperature and potential
evaporation decrease with elevation (Table 1). The
rainfall pattern is less clearly defined. Although
80–95% of the rain falls from May to November, there
is a gradient of increasing rainfall from east to west –
southwest in the lowland Llanos. Rainfall decreases with
elevation except in the Gran Sabana (Table 1). The
length of the dry season varies from 2–3 months in the
highlands, to 5–6 months in the lowlands.
The Trachypogon savannas overlay diverse sub-
strates varying from creasts and midslopes of the
Coastal Mountains, recent alluvial and aeolian plains
in the lowland Llanos and old sandstones of the
Precambrian Guiana Plateau (Cole, 1986). The soils
are mostly Ultisols or Oxisols which are acidic,
oligotrophic and low in organic matter. Soil texture
varies from clayey to almost pure sand which influences
their water and nutrient retention capacity. Fertility
tends to be higher in the highlands and lowest in the
sandy soils. A superficial lithoplinthic crust (known
locally as ‘‘arrecife’’ or ‘‘ripio’’) is conspicuous in several
sites.
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Z. Baruch / Flora 200 (2005) 49–64 51

Table 1. List of sampled sites, the numbers identify the site in Fig. 1

Savanna type

Type I Type II Type III Type IV

Number and name of 10 4 17 6

sampled sites
El Volcán (1), Caricuao Aluepa (11), Iworibo Calabozo (15), Icaros La Iguana (32), El Tigre
(2), Magdalena (3), La (12), Yuruanı́ (13), Km (16), L. de Hierro (17), (33), San Joaquı́n (34),
Encantada (4), 217 (14) Guasipati (18), Los Pijiguaos (35),
Araguita (5), La Mora Mochima (19), S.Elena Médanos (36), Puerto
(6), La Victoria+660 (20), Uverito (21), S. Paez (37)
(7), La Victoria (8), Las Barbara (22), La Isla
Lapas (9), Petaquire (23), Barinas (24),
(10) Mataclara (25), El Pao
(26), S. Carlos (27),
Pijiguaos (28), Caicara
(29), P. Ayacucho (30),
S. Rita (31)
Elevation range (m) (1100–1980) (1130–1300) (10–800) (30–175)
Air temperature range (16.8–25.6) 22.6 (21.4–28.2) (25.9–28.3)
(1C)
Rainfall range (mm) (762–1100) 1851 (700–2238) (1027–1935)
Pan evaporation range (1060–1700) 2100 (1940–2400) (1946–2699)
(mm)
Humidity index range (0.45–0.90) 0.93 (0.28–0.93) (0.38–0.81)
(HI)
Soil P range (ppm) (5.0–10.0) (9.0–11.0) (4.0–14.0) (5.0–7.0)
Soil K range (ppm) (50.0–300.0) (8.0–88.0) (8.0–188.0) (8.0–24.0)
Soil Ca range (ppm) (54–595) (115–255) (95–475) (95–215)
CEC range (8.2–20.0) (2.1–12.6) (1.1–16.8) (0.1–4.5)
(meq 100 g1)
Fertility index range (53.3–189.9) (89.6–113.4) (47.7–157.1) (47.7–78.3)
(FI)
Sand range (%) (39.6–63.6) (73.6–87.6) (45.6–89.6) (74.6–91.6)
Org. matter range (%) (2.7–6.5) (1.5–4.1) (0.7–5.0) (0.6–1.9)
pH range (4.8–7.6) (4.1–4.7) (4.2–6.0) (4.4–4.7)
Lithoplintite () (+) & () (+) & () ()

Range of elevation, climate and soil characteristics of the study sites grouped into the four savanna types. HI and FI are humidity and fertility
indices, respectively. The presence of local soil lithoplintic hardpan is shown as (+) or ().

All surveyed savannas, except some Gran Sabana
sites (which are in a National Park) and the fenced
Calabozo site in the Llanos, are grazed with varying
intensity. Fires occur each 1–3 years during the dry
season, mainly in the Llanos and Gran Sabana. Except
in the Gran Sabana, African grasses have spread and
displaced native plants, creating dense and almost
monospecific stands (Baruch, 1996; Williams and
Baruch, 2000).
Methods
Vegetation was surveyed in all large savanna geo-
morphological regions: spaced from east to west along
the central Llanos, in the Coastal Mountains and in
areas that have not been analyzed floristically before
such as the Gran Sabana and the south margin of the
Orinoco. Specific sites were selected by the following
criteria: (a) T. plumosus or T. vestitus as the dominant
grasses; (b) savannas not subjected to flooding; (c) lack
of evidence of very recent (1 year) fires; and, (d) at least
500 meters from roads or human occupation. Field
work was completed during the rainy seasons of 2 years.
In total, 293 species of graminoids, herbs and shrubs
were censused (Appendix A). Nomenclature follows
Aristeguieta (1966), Vareschi (1970), Ramia (1974), and
Steyermark and Huber (1978). Voucher samples are
deposited in the herbarium of Universidad Simón
Bolı́var.
Sampling was done in 376 plots from 37 sites (Fig. 1
and Table 1). At each site, a 50–100 m long transect was
laid out and 5–20 one m2 plots were randomly located
 ARTICLE IN PRESS
52 Z. Baruch / Flora 200 (2005) 49–64
along it with a quadrat frame sub-divided into
10  10 cm segments. In each plot plant cover was
measured by counting the contacts with a pin lowered
100 times at a regular pattern. Density was determined
by carefully delimiting and counting individuals of each
species. The area of bare soil was determined and total
plant cover was calculated. The importance value index
(IVI) was obtained for each species as the sum of the
relative cover (RC) and relative density (RD) (Mueller-
Dombois and Ellenberg, 1974). Pooling of IVI values
was done for all plots within a site. Concurrently, 3 soil
samples (0–20 cm depth) were taken and pooled from
each site. Soil texture was determined by the Bouyoucos
hydrometer method, available phosphorus was deter-
mined by the molybdic-blue method (Murphy and
Riley, 1963) and available potassium and calcium were
determined by flame spectrophotometry. Cation ex-
change capacity (CEC) was determined by extraction
with NH4Cl solution followed by spectrophotometry. A
synthetic fertility index (FI) was calculated as the sum of
the relative values (with respect to their maximum) of K,
Ca and P for each site. The maximum value of FI is 300.
Organic matter content, by the Walkley and Black’s
method (Jackson, 1982), and pH (1:2.5 in water) were
also determined. Mean air temperature, rainfall and pan
or potential evaporation data from 27 climatological
stations, maintained by the Venezuelan government
(Ministerio del Ambiente and Air Force), near the study
sites were obtained. A synthetic humidity index (HI) was
calculated as the ratio between average annual rainfall
and pan evaporation. The number of dry months, either
2–3 or 4–6 was considered as a binary environmental
variable. Disturbance, as recent evidences of fire and
grazing, was estimated visually and scaled from low to
high from 1 to 3. This scale combines magnitude and
frequency of disturbances. Other qualitative variables,
as the presence of the lithoplintic crust on the soil and of
woody plants were included for comparison.
Data analysis was performed on the species IVI
averaging all plots per site. All numerical analyses were
done with the PC-ORD, V. 4 package (McCune and
Mefford, 1999). Species with less than 2 entries were
deleted to increase the definition of the results. For each
savanna type, total and mean (per site) species richness
were obtained. Beta-diversity was calculated as the ratio
of total to mean species richness. Data were subjected to
cluster analysis using Euclidean distance and the Ward
group linkage method. In addition, to identify species
with particular diagnostic value and to confirm cluster-
ing results, the floristic data were classified with the two-
way indicator species analysis (TWINSPAN) (Hill,
1979).
The floristic data and environmentally important
variables were subjected to canonical correspondence
analysis (CCA) (Jongman et al., 1987; ter Braak, 1987)
to relate the internal structure of the data matrix with
ecologically meaningful environmental gradients. The
regression coefficients of the environmental variables
generated by the analysis and the total significance of
the ordination were tested with a Monte-Carlo permu-
tation test under the null model.
Results
Although clustering is an agglomerative classification
technique and TWINSPAN is divisive, both produced
comparable results. In addition, TWINSPAN provided
indicator species. Integrating the results of both
classifications methods with geographic affinities allows
for an ecologically sound scheme of 4 major savanna
types. The first division of the clustering dendrogram
generated a group of sites from the sandy and treeless
savannas (Type IV) where the indicator and dominant
grass is T. vestitus (Fig. 3). The third division generated
an homogeneous cluster with the sites above 700 m in
the Coastal Mountains (Type I) (Figs. 2 and 3). This
group was also generated by the first division in
TWINSPAN characterized by the indicator species
Rhynchospora rugosa and Achyrocline satureioides. The
cluster of four sites from the isolated Gran Sabana was
generated by the fourth division in the dendrogram
(Type II) (Fig. 3) and by the second division of
TWINSPAN with the presence of indicator species as
Rhynchospora barbata, Axonopus anceps and Lagen-
ocarpus rigidus. The majority of the lowland sites (Type
III) did not conform an homogeneous cluster but were
scattered in the two remaining groups formed after the
second and the third division (Fig. 3).
Ten sites from the Coastal Mountains conform the
Type I savannas (Table 1 and Fig. 2, bottom). They are
relatively cool, humid and fertile, contain the largest
number of species and the highest percentage of
exclusive species (Table 2). Type I savannas differ from
the rest by the presence and importance of the
melastome shrubs of the genera Miconia and Clidemia.
These are joined by the graminoids Andropogon
condensatus, A. tener, Eragrostis fragilis, Panicum
aciculare, Rhynchospora rugosa, Setaria geniculata and
Scleria hirtella, herbs as Achyrocline satureioides,
Buchnera rosea, Coccocypselum lanceolatum, Chaptalia
runcinata, Eupatorium amygdalinum, Gnaphalium pur-
pureum, Ichthyothere terminalis, Orthopappus angustifo-
lius, Polygala monticola, Pterocaulon virgatum,
Sauvagesia sprengelii, Schoenocaulon officinale, Stevia
elongata and Trimezia martinicensis, and other shrubs as
Bejaria glauca, Gaylussacia sp., Hypericum caracasanum,
Hyptis sp., Myrica caracasana, Psidium guajava and
Vaccinium sp. among the total 194 species censused
(Appendix). In this type, the sub-dominant grass is
Axonopus pulcher. Here, the invader African grass
 ARTICLE IN PRESS
Z. Baruch / Flora 200 (2005) 49–64
Fig. 2. Plate showing three savanna types. Top: Woody
savanna of Type III in the Calabozo site (15). Middle: treeless
savanna of Type II in the Aluepa site (11). Bottom: shrubby
savanna of Type I at intermediate elevations in the Las Lapas
site (9).
Melinis minutiflora forms isolated but relatively large
patches displacing the native savanna community.
Type II savannas, with 4 sites from the treeless Gran
Sabana (Table 1 and Fig. 2, middle) are relatively
isolated from the rest by distance and elevation. They
scored second in total species richness (Table 2) and
have some typical dominant and endemic species from
the Guianan region (Berry et al., 1995) such as the
graminoids Axonopus anceps, Echinolaena inflexa, La-
genocarpus rigidus, Panicum chnoodes, Paspalum con-
tractum, P. stellatum, Rhynchospora barbata, Scleria
53
cyperina, and herbs such as Buchnera palustris, Polygala
hygrophila, Radiella esenbeckii, Siphanthera cordifolia,
and Sisyrinchium vaginatum. Leguminous plants are
noticeably absent from this savanna type whereas
termitaria are conspicuous in some sites.
Type III savannas form a large heterogeneous group
as evidenced by its relatively high b-diversity and area
covered (Tables 1 and 2; Figs. 1 and 2, top). It represents
the core of the lowland Llanos stretching out from east
to west north of the Orinoco and, in some places, also
south of the river. This is the typical savanna landscape
with dispersed small trees, mainly Curatella americana,
Byrsonima crassifolia and Bowdichia virgilioides. Axo-
nopus canescens is the sub-dominant grass in most of the
sites and other important graminoids are Andropogon
fastigiatus, Bulbostylis spp., Leptocoryphium lanatum,
Dichromena ciliata, Rhynchospora albo-marginata, Schi-
zachyrium sanguineum, and Sporobolus cubensis. Le-
gume herbs are numerous and an important component
of the vegetation, as Aeschynomene brasiliensis, Clitoria
guyanensis, Eriosema crinitum, Galactia jussieuana,
Phaseolus linearis, and Zornia reticulata. Other impor-
tant herbs are Borreria capitata, Evolvulus sericeus,
Merremia aturensis, and several species of Polygala. This
savanna type includes patches of lithoplintite outcrops
covered by an low cover and species poor herbaceous
community (Velásquez, 1965). In the most fertile sites,
the African invader grass Hyparrhenia rufa has dis-
placed native communities.
Type IV savannas (6 sites) occupy sandy unfertile soils
on both sides of the Orinoco river and show a strong
dominance by T. vestitus. Here, plant cover and species
richness and diversity are the lowest and woody plants
overtopping the grass stratum are absent (Table 2).
The first two axes of the ordination by CCA (Fig. 4)
accounted for 15.6% of total variation. This seems
ecologically realistic for an hetereogeneous group of
sites over a large area. The Monte-Carlo test indicates
statistically significant eigenvalues for the three ordina-
tion axes (for all, po0:005). The relationship of selected
environmental variables with the first ordination axis
was strongly significant (p ¼ 0:002) and less so
(p ¼ 0:08) for the second axis as indicated by the
Monte-Carlo randomization test. The CCA biplot
shows a clear separation between the highland savannas
in the Coastal Mountains (Type I) and the rest of the
sites which confirms classification results (Fig. 4). These
sites are correlated with relatively high fertility (high
CEC), water availability (HI) and short dry seasons
(Fig. 4). All sites from savanna Type II and four from
Type IV form relatively isolated clusters defined by low
fertility and high rainfall (Fig. 4). Although rainfall is
high in both groups, HI separates them on axis 2
because Type IV, with higher average pan evaporation,
has lower HI than Type II (Fig. 4 and Table 1). The
Type IV savannas correlate strongly with high soil sand
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54 Z. Baruch / Flora 200 (2005) 49–64

Fig. 3. Dendrogram of the cluster grouping of the study sites. Grouping was performed using Euclidean distance and the Ward
method. Species with less than 2 entries in the matrix were deleted from the analysis. Symbols correspond to the savanna types
according to Fig 1 and Table 1.

Table 2. Vegetation and floristic characteristics of the four savanna types

Savanna type

Type I Type II Type III Type IV

Number of sites 10 4 17 6
Total species richness 194 59 85 53
Mean species richness (per site) 38.9716.4 13.871.7 17.878.5 14.574.4
b-diversity 4.98 3.55 4.77 3.65
Species limited to type 131 16 40 16
% of Total spp. richness 67.5 27.1 47.0 30.2
% Cover, mean and range 93.8 (100.0–88.2) 80.9 (87.9–74.6) 82.5 (98.3–61.3) 79.1 (87.8–71.1)
Woody elements (S) (TL) (T) & (TL) (TL)

b-Diversity was calculated as total species richness/mean species richness. Plant cover is shown as mean and range. The presence of woody elements
in the savanna is designed as (S) ¼ shrubs, (T) trees and (TL) treeless.

content and low percentage plant cover (Fig. 4). Most of
the numerous and heterogeneous sites of Type III, were
placed in the remaining quadrant of the biplot defined
by low water availability and fertility (Fig. 4) where they
overlap with the remaining two sites from Type IV.
Discussion
Although all savannas had a strong dominance by
Trachypogon species, they showed substantial structural
and floristic heterogeneity and a complex relationship
with the local environment. Classification and ordina-
tion methods generated results that were complementary
and ecologically sound. The results of both methods
indicated that elevation was a main determining
environmental factor which clearly separated the rela-
tively species rich, fertile, moist and cool savannas of the
Coastal Mountains (Type I) from the rest. Water and
nutrient availability, caused by climatic or edaphic
factors, were found to be the determining environmental
variables in savanna demarcation either by species
richness, community composition or physiognomy. This
 ARTICLE IN PRESS
Z. Baruch / Flora 200 (2005) 49–64 55

UVER

SJOAQ MOCHI

ELPAO SCARL
IGUAN
MATCL VC66
SBARB
GUASI
BARIN
PIJIG
CAICA
LAISL
MAGDA
ARAG
ICAR
CALAB
PIJ40
SRITA CEC PETA

ELTIG LOMAH VICT ENCAN

PPAEZ % SAND
VOLCA
MEDAN

RAINFALL STELE HI ELEV. CARIC

MORA

PAYAC LAPA

YURUA

Type I
Type II
Type III
Type IV
KM217
IWORI
ALUEP

Fig. 4. CCA biplot of the study sites and main environmental variables. Species with less than 2 entries in the matrix were deleted
from the analysis. Symbols correspond to the savanna types according to Fig 1 and Table 1. Names are the first 4–5 letters of the
sites names in Table 1.

relationship confirms, at the regional level analyzed
here, their ecological importance as the main limiting
factors in tropical savannas (Frost et al., 1986; Medina
and Silva, 1990). In response to these limiting factors, T.
plumosus, and probably other savanna plants, show
several resource conservation traits such as stomatal
closure, osmotic adjustment, high leaf mass area ratio,
and low leaf nutrient content (Baruch and Fernández,
1993; Baruch and Gómez, 1996).
In addition to climatically defined water availability,
the length of the rainless period is also important in
defining this seasonal ecosystems. It separated the
highland Type I sites, with 2–4 rainless months, from
the other savanna types with 5–6 dry months each. San
José et al. (1998) found that the length of the dry season
was an important variable in the analysis of Venezuelan
and Colombian lowland savannas. Soil texture was
correlated with site ordination and defines the capacity
of soils to retain nutrient and water. The sandy soils of
Types II and IV savannas have a water holding capacity
of 3.1% (water content at field capacity minus water
content at wilting point) whereas some Type I savannas
retain 16.5% (Z. Baruch, unpublished data). At a very
local level, Sarmiento and Monasterio (1969) used soil
texture to distinguish communities dominated by T.
plumosus from those dominated by T. vestitus.
Soil fertility was also significantly correlated with site
ordination. Soil organic matter was correlated positively
with nutrient availability whereas soil sand content was
negatively correlated. Here, nutrient availability was
quantified by the FI or Cation Exchange Capacity.
Highland soils (Type I) with a high organic matter
 ARTICLE IN PRESS
56 Z. Baruch / Flora 200 (2005) 49–64
content and CEC were more fertile than the sandier soils
from Savanna Types II and IV. This limits plant growth
in the later savanna types (Baruch and Gómez, 1996).
Similar overriding influence of nutrient availability on
the composition and structure of tropical savannas has
been reported in other studies from Venezuela and
Colombia (Beard, 1953; San José et al., 1998; Susach,
1989) and from Guyana, Surinam and Brazil (Good-
land, 1966; Goodland and Pollard, 1973; Goldsmith,
1974; Lopes and Cox, 1977; Miranda et al., 2003; van
Donselaar, 1965). In addition, it has been proposed that
fertility edaphic gradients were the main cause for the
vegetation gradual transformation from grassland or
treeless savanna to the typical woody savanna (Good-
land and Pollard, 1973; Lopes and Cox, 1977; Sarmiento
and Monasterio, 1969).
Fire is another of savanna’s major characteristics
(Frost et al., 1986) and has been invoked as one of the
factors determining its structure and composition
(Beard, 1953; Blydenstein, 1962). Here, fire and grazing
disturbances had no weight in defining the ordination
results. When considered jointly, these two disturbances
balance even among the savanna types. Due to their
higher humidity and shorter dry season, Type I
savannas burn less frequently but grazing is relatively
important. In contrast, the other savanna types burn
almost annually and grazing is important only in Types
III and IV. However, the role of fire is strongly
evidenced when savannas are protected. The woody
component increases and the relative importance of
several grass species changes in Venezuelan (San José
and Fariñas, 1991) and in Brazilian (Moreira, 2000)
savannas.
The presence and prominence of woody elements,
determines savanna physiognomy and landscape. It is a
partial result of the regional differences in biophysical
variables and aids in the interpretation and classification
of the savannas. The conspicuous shrubs (1–2 m tall) of
the highland Type I savannas are consequence of their
favorable water and fertility status and reduced fire
frequency. They also suggest that these savannas are a
successional stage towards the original forest (Tamayo,
1943; Vareschi, 1970). Dispersed trees in the lowland
Type III savannas point to relatively deep soils with a
reasonable water and nutrient storing capacity (Var-
eschi, 1992; Walter, 1973). Their absence or low
prominence (as they grow dwarfed without overtopping
the herbaceous matrix) from Types II and IV savannas
is probably due to a combination of low water retention
and fertility caused by sandy soils and to frequent fires.
Total species richness differed notably among savan-
na types as consequence of the interaction of the
regional environment and the available species pool.
Type I savannas from the Coastal Mountains grade
continuously from 700 to 1900 m asl. Here, mean plant
cover and species number are the highest. Total species
richness is also the highest and 67% of the species are
exclusive of this site (Table 2). The richness of this flora
is caused by the blend of typical lowland savanna
elements and those of the sub-páramo, which become
more prominent as elevation increases (Steyermark and
Huber, 1978; Vareschi, 1970). However, where the
African invader grass Melinis minutiflora has displaced
the community, species richness decreases dramatically
(Baruch, 1996). The Gran Sabana (Type II) has an
exclusive assemblage of species accompanying T.
plumosus (Huber, 1995). This is probably caused by
the unusual combination of relatively cool temperature
(mean 22.6 1C), high rainfall and HI, low soil fertility
and the influence of elements of the Guiana flora. An
interesting feature of these savannas is the absence of
legumes, so conspicuous in other savannas.
Type III savannas are the most extended and could be
considered the ‘‘typical’’ savanna landscape. However,
herbaceous species richness and plant cover are rela-
tively low. They decrease even further on the lithoplintic
soil or ‘‘ripio’’ patches which, in some sites, forms a
mosaic within the deeper soils of the savanna. The
particularity of the ripio patches is probably due to the
lower soil water retention capacity and to the impedance
to root growth (Pieters and Baruch, 1997) which
confirms the results of Velásquez (1965) and San José
et al. (1998). The sandy Type IV savannas on both
banks of the Orinoco form a mosaic of apparently
‘‘treeless’’ savannas with low plant cover probably
caused by increased competition for the scarse soil
water and nutrients. Here, the densely pubescent T.
vestitus replaces T. plumosus as the dominant species
and small stunted trees do not overtop the grasses.
The savanna classification presented here is partially
consequence of the regional geomorphology. Type I
savannas, on crests and mid-slopes of the Coastal
Mountains, have soils that rejuvenate gradually by
erosion thus maintaining fertility. The sandstones of the
Precambrian Roraima Formation underlay the extre-
mely nutrient poor Type II savannas. The Type III
savannas are either on alluvial Quaternary terraces in
the west of the country or on the Pliocene–Pleistocene
Mesas that underlay the central and eastern Llanos.
Finally, aeolian plains underlay the Type IV savannas
(Cole, 1986; Sarmiento, 1983).
This study shows that the composition and structure
of the Trachypogon savannas are related to the specific
water and nutrient availability of the major geomor-
phological regions of Venezuela. Each of the previous
studies found similar results for their particular savan-
nas sites. Blydenstein (1962), Velásquez (1965), Sar-
miento and Monasterio (1969) and Ramia (1993)
described the non-flooded central Llanos and found
that the dominance of Trachypogon species and the
importance of woody elements depend on edaphic
factors such as texture, depth of water table and
 ARTICLE IN PRESS
Z. Baruch / Flora 200 (2005) 49–64
nutrient content. Susach (1989) analyzed a region of the
Central Llanos close to the Orinoco river. The non-
flooded savannas of this area correspond to our site
Santa Rita (31). Using the nomenclature of the Zürich –
Montpellier school, he classified the vegetation of the
non-flooded savannas into the order Trachypogonetalia
plumosi belonging to the class Leptocoryphio-Trachy-
pogonetea, similar to those savannas described by van
Donselaar (1965) in the Guyanas. In their analysis, San
José et al. (1998) included the Colombian Llanos. They
classified the savannas into three types also based on soil
fertility and water availability. Huber (1995) described
the Gran Sabana vegetation as sub-mesothermic savan-
nas with a large influence of the Guianan flora. Vareschi
(1970) described the savannas in the Coastal Mountains
and found co-dominance by T. plumosus and Axonopus
pulcher. The Trachypogon dominated savannas extend
to the Guianas and the Roraima region in Brazil and
were grouped in the class Leptocoryphio-Trachypogo-
netea (van Donselaar, 1965). Goodland (1966) analyzed
the vegetation of the Rupununi-Rio Branco region and
found that soil moisture and nutrients were the
dominant environmental variables controlling vegeta-
tion distribution. Floristics and physiognomy were
similar to those of the Calabozo site (Type III
savannas). Although many species differ, correspon-
dence in savanna physiognomy can be readily estab-
lished between our types and those found in the
Brazilian cerrado. Type I shrubby savanna resembles
the ‘‘campo sujo’’, the Types II and IV treeless savannas
correspond to the ‘‘campo limpo’’ whereas the extended
Type III savanna corresponds to the ‘‘campo cerrado’’
(Oliveira-Filho and Ratter 2002; Sarmiento, 1983).
Savanna stability, in terms of structure and composi-
tion of their native communities, could be easily
disrupted by the increasing use of this ecosystem. Run-
off from fertilized crops and pastures might increase the
nutrient availability in the native savanna and promote
changes as increased plant cover and/or species replace-
ment through invasion of exotic grasses (Barger et al.,
Appendix A
Species list from the four Trachypogon savanna types. Species marked with xx designate those with IVI41
Savanna types
Acanthaceae
Ruellia geminiflora H. B. K.
Amaryllidaceae
Hypoxis decumbens L.
Asteraceae
Achyrocline satureioides DC.
Ageratum conyzoides L.
Chaptalia nutans (L.) Polak
C. runcinata H. B. K.
57
2003; Baruch, 1996; Parsons, 1972; Pieters and Baruch,
1997; Williams and Baruch, 2000). The alteration of the
fire regime may lead to either species loss or the
encroachment of the woody component (Klink et al.,
1993; San José and Fariñas, 1991; Silva, 1996). The most
fragile savannas appear to be Types II and IV due to
their extremely nutrient poor sandy soils. In addition,
the particular climatic–edaphic blend of the Type II
(Gran Sabana) supplemented to its isolation, signals its
vulnerability. This analysis of the vegetation–environ-
ment relations and the classification of the Venezuelan
savannas is also relevant for the savannas in Colombia,
Brazil and Guyana and provides a base line for other
studies intended to conserve and restore this ecosystem.
Acknowledgments
Universidad Simón Bolı́var supported this study
through the Decanato de Investigaciones. I wish to
thank S. Baruch, T. da Silva, M. Edreira, A. Gols, M.
Rodriguez , and V. Salas who helped in data collection
and analysis. Sociedad Venezolana de Ciencias Natur-
ales and Universidad Simón Rodriguez provided per-
mits and lodging at their field stations, Biológica de los
Llanos and La Iguana, respectively. Ministerio del
Ambiente (MARNR) provided some of the climatic
data through the collaboration of Ing. H. Fonseca. The
Instituto Nacional de Parques (INPARQUES) granted
permission to collect in the Avila, Mochima and Gran
Sabana National Parks. Botanical samples were gener-
ously identified by M. Ramia, F. Delascio, O. Huber
and D. Velásquez. I wish to thank O. Solbrig and G.
Romero for their hospitality and logistic support at
Harvard University Herbaria where the first version of
this paper was written and to R.B. Jackson and the
Jackson’s Lab group of Duke University for hospitality
and valuable comments.
I II III IV
x x
x
xx
x
x
xx
 ARTICLE IN PRESS
58 Z. Baruch / Flora 200 (2005) 49–64

Savanna types I II III IV

Clibadium sp. x
Conyza chilensis Spreng. x
Erechtites hieraciifolia (L.) Raf. ex DC. x
Eupatorium amygdalinum Lam. x x
Gnaphalium purpureum L. x
G.simplicicaule Willd. ex Spreng. x
Ichthyothere terminalis (Spreng.) Blake x xx x
Ichthyothere sp. x
Orthopappus angustifolius (Sw.) Gleason x x
Pectis carthusianorum Less. x
P. ciliaris L. x
P. elongata H. B. K. x
Praxelis pauciflora R.M.King & H.Rob. x
Pterocaulon virgatum (L.) DC. x
Stevia elongata H. B. K. xx
S. lucida Lag. x
S. wageneri Hieron. x
Wedelia caracasana DC. x
Bignoniaceae
Jacaranda obtusifolia Bonpl. x
Boraginaceae
Heliotropium polyphyllum Lehm. x
H. angiospermum Murray x
Burmanniaceae
Alteria aphylla Barnhart x
Burmannia capitata Mart. x
Caesalpiniaceae
Cassia biflora L. x
C. calycioides (DC.) Greene x
C. cultrifolia H. B. K. x
C. diphylla L. x
C. flexuosa L. x x
C. hispidula Vahl. x x
C. patellaria DC. x x
C. rotundifolia Pers. x
C. serpens L. x
Cassia sp. x x x
Caryophyllaceae
Polycarpaea corymbosa Lam. x x
Convolvulaceae
Evolvulus argenteus Pursh x
E. elegans Moric. x
E. filipes Mart. x
E. pterocaulon Moric. x
E. sericeus Sw. x x
Ipomoea asarifolia Roem. & Schult. x
I. capillacea Gray x x
Jacquemontia sphaerostigma Rusby x
Merremia aturensis (H. B. K.) Hall. f x
Cyperaceae
Bulbostylis capillaris (L.) Kunth x xx xx xx
B. conifera Kunth x xx xx xx
B. junciformis (H. B. K.) Lindman xx x
 ARTICLE IN PRESS
Z. Baruch / Flora 200 (2005) 49–64 59

Savanna types I II III IV

B. paradoxa (Speng.) Lindman x xx xx

Cyperus sp. x
C. flavus (Vahl.) Nees x
Dichromena ciliata Vahl xx xx x
Fimbristylis sp. x
Lagenocarpus rigidus Nees xx
Rhynchospora albo-marginata Kük. xx
R. barbata (Vahl) Kunth xx x
R. caracasana (Kunth) Boeckl. x
R. duckei R. Gross x
R. globosa (H.B.K.) Roem. et Schult. xx xx xx
R. rugosa (Vahl) Gale xx
R. subplumosa C.B.Clarke x x
Scleria bracteata Cav. x
S. cyperina Kunth xx x
S. distans Poir. x
S. hirtella Sw. xx
S. microcarpa Nees x
Dilleniaceae
Curatella americana L. xx x
Ericaceae
Befaria glauca Bonpl. x
Gaylussacia sp. x
Euphorbiaceae
Croton hirtus L’Herit. x
Euphorbia dioica Hieron. x
E. glomerifera (Millsp.) L. C. x
Phyllanthus lathyroides H. B. K. x
Sebastiania corniculata Pax x
Flacourtiaceae
Casearia sylvestris Sw. x
Gentianaceae
Schultesia benthamiana Klotzsch x x
S. brachyptera Cham. x x
Haemodoraceae
Schieckia orinocensis Meissn. x x
Hypericaceae
Hypericum caracasanum H. B. K. x
Iridaceae
Cypella linearis (H.B.K) Baker x
Sisyrinchium alatum Hook. x
S. vaginatum Spreng. x
S. micranthum Cav. x
Trimezia martinicensis (Jacq.) Herb. x
Lamiaceae
Hyptis atrorubens Briq. x
H. brachiata Briq. x x
H. divaricata Pohl ex Benth. x
H. fulva Epling x
H. lantanifolia Poit. x
H. mutabilis (Rich.) Briq. x
H. pulegioides Briq. x
Hyptis sp. x
 ARTICLE IN PRESS
60 Z. Baruch / Flora 200 (2005) 49–64

Savanna types I II III IV

Lentibulariaceae
Utricularia amethystina Salzm. ex A.St Hil. & Girard x
Liliaceae
Schoenocaulon officinale (Schltdl. & Cham.) A.Gray ex Benth. xx
Lythraceae
Cuphea antisyphilitica Benth. x
C. micrantha H. B. K. x
Malvaceae
Cienfuegosia affinis (H. B. K.) Hochr. x
Hibiscus furcellatus Lam. x
Peltaea speciosa (H. B. K.) Standl. x
Sida linifolia Juss. ex Cav. x
Sida rhombifolia L. x
Melastomataceae
Clidemia rubra Mart. x
C. sericea D. Don x x
Comolia veronicaefolia Triana x
Miconia albicans (Sw.)Triana x
M. ciliata (L. C. Rich.) DC. x
M. rubiginosa (Bonpl.) DC. x
Siphanthera cordifolia Gleason x
Tibouchina longifolia (Vahl.) Baillon x
Mimosaceae
Mimosa martensis L. x x
M. pudica L. x
Mimosa sp. x
Myricaceae
Myrica caracasana H. B. K. x
Myrtaceae
Eugenia pauciflora DC. x
Psidium guajava L. x
P. salutare Berg x
Ochnaceae
Sauvagesia sprengelii A. St Hil. x
Orchidaceae
Bletia lansbergii Rchb.f. x
Cyrtopodium parviflorum Lindl. x
Habenaria sp. x
Orchidaceae sp. x
Papilionaceae
Aeschynomene brasiliensis (Poir.) DC. x xx
A. elegans Schltdl. & Cham. x x
A. paniculata Willd. ex Vogt x
Bowdichia virgiloides H.B.K. xx
Calopogonium mucunoides Desv. x
Centrosema venosum Mart. ex Benth. x
Clitoria guyanensis (Aubl.) Benth. x xx x
C. rubiginosa (A. Juss.) Pers. x
Crotalaria stipularis Desv. x x x
Desmodium barbatum (L.) Benth. x x
D. campyloclados Hemsl. x
D. canum Schinz & Thellung x
D. incanum DC. x
 ARTICLE IN PRESS
Z. Baruch / Flora 200 (2005) 49–64 61

Savanna types I II III IV

D. pachyrrhizum Vogel x
D. procumbens (Mill.) Hitchc. x
D. scorpiurus (Sw.) Desv. x
D. sericophyllum Schltdl. x
Eriosema crinitum (H. B. K.) G. Don x xx xx
E. simplicifolium (H.B.K.) G. Don x
Galactia jussieuana Kunth xx x
Indigofera lespedezioides H. B. K. x
I. pascuorum Benth. xx x
Macroptilium gracile Urb. x
Phaseolus diversifolius Pers. x
P. erythroloma Mart. ex Benth. x
P. gracilis Poeppig ex Benth. x
P. lathyroides L. x
P. linearis H. B. K. x xx xx
Rhynchosia edulis Griseb. x
Stylosanthes capitata Vog. x
S. guianensis (Aubl.) Sw. x x x
S. humilis H. B. K. x x
S. sericea S. F. Blake x
Tephrosia leptostachya DC. x xx
T. tenella A. Gray x
Zornia curvata Mohlenbrock x
Z. diphylla Pers. x x
Z. filifoliola Domin x
Z. reticulata Sm. x
Zornia sp. x
Poaceae
Andropogon angustatus (Presl) Steud. x
A. bicornis L. x x
A. condensatus H. B. K xx xx
A. fastigiatus Sw x
A. selloanus (Hack.)Hack. x xx
A. semiberbis (Nees) Kunth x x
Andropogon sp. x
A. tener (Nees)Kunth xx
Aristida recurvata H. B. K. x x x
A. torta Kunth xx x
Arundinella confinis (Schult.) Hitchc. & Chase xx
Axonopus anceps (Mez.) Hitchc. xx x
A. canescens (Nees & Trin.) Pilger x xx xx xx
A. chrysoblepharis (Lag.) Chase x
A. pulcher (Nees) Kuhlmann xx xx
A. purpusii (Mez.) Chase xx x
Axonopus sp. x xx x x
Digitaria fragilis (Steud.) Luces x
D. villiculmis Henr. x
Echinolaena inflexa (Poir.) Chase xx x
Elionurus planifolius Renvoize x
E. tripsacoides (H. B. K.) Hack. x
Eragrostis airoides Nees xx
E. fragilis Swallen x
E. polytricha Nees x
 ARTICLE IN PRESS
62 Z. Baruch / Flora 200 (2005) 49–64

Savanna types I II III IV

Gymnopogon foliosus Nees xx

Hackelochloa granularis (L.) Kuntze x
Hyparrhenia rufa (Nees) Stapf x x
Leptocoryphium lanatum (H. B. K. ) Nees x x
Melinis minutiflora Beauv. xx
Mesosetum loliiforme Hitchc. xx
M. chaseae Luces xx
Panicum aciculare Desv. ex Poir. x
P. chnoodes Trin. xx
P. micranthum H. B. K. x xx
P. olyroides H. B. K. x
P. orinocense Willd. ex Spreng. x
P. polycomum Trin. xx x
Panicum sp. x
P. sphaerocarpon Ell. x
P. tenuissimum Benth. x
Paspalum contractum Pilger xx
P. carinatum Humb. & Bonpl. ex Flugge x
P. gardnerianum Nees x
P. pilosum Lam. x
P. plicatulum Michx. x
Paspalum sp. x
P. stellatum Humb. & Bonpl. x xx
Radiella esenbeckii (Steud.) Cald. & Soderstr. x
Schizachyrium tenerum Nees xx
S. sanguineum (Retz.) Alston xx xx
Setaria geniculata (Lam.) Beauv. xx
Sporobolus cubensis Hitchc. x xx xx x
Trachypogon plumosus (H. & B.) Nees xx xx xx xx
T. vestitus Anders xx xx
Thrasya petrosa (Trin.) Chase x
Polygalaceae
Polygala diversifolia L. x
P. fendleri Chodat x
P. hygrophila H. B. K x
P. leptocaulis Torr. & Gray x
P. longicaulis H. B. K. x
P. monticola H. B. K. x x
P. paniculata L. x
P. savannorum Chodat x
Polygala sp. x
P. variabilis H. B. K. x xx
Polygonaceae
Coccoloba caracasana Meissn. x
Pteridophyta
Anemia villosa H. & B. x
Pteridium caudatum (L.) Maxon xx
Rubiaceae
Borreria capitata (Ruiz & Pavon) DC. x xx xx
B. ocimoides (Burm. f.) DC. x
B. verticillata (L.) G.F.W.Meyer x
Coccocypselum lanceolatum (Ruiz & Pavon) Pers. x
Declieuxia fruticosa (Willd.) Kuntze x x x
 ARTICLE IN PRESS
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Savanna types I II III IV

Diodia teres (Sw.) Steyerm. x x x

Palicourea rigida H. B. K. x
Relbunium hypocarpicum (L.) Hemsl. x
Roupala complicata H. B. K. x
R. montana Aubl. x
Sipanea hispida Benth. ex Wernh. x
Sipanea sp. x
Sapindaceae
Dodonaea viscosa Jacq. x
Scrophulariaceae
Buchnera longifolia H. B. K. x
B. palustris Aublet x x
B. rosea H. B. K. x
B. weberbaueri Diels x x x
Sterculiaceae
Waltheria americana L. x
Turneraceae
Piriqueta cistoides (L.) Griseb. x
Turnera ulmifolia L. x x
Verbenaceae
Lantana camara L. x
Lantana caracasana Turcz. x
Xyridaceae
Xyris garcia-barrigae Idrobo & L. B. Smith xx x

References Blydenstein, J., 1962. La sabana de Trachypogon del Alto Llano

Aristeguieta, L., 1966. Flórula de la Estación Biológica de los
Llanos. Bol. Soc. Venez. Cien. Nat. 110, 228–307.
Barger, N.N., D’Antonio, C.M., Ghneim, T., Cuevas, E.,
2003. Constraints to colonization and growth of the
African grass Melinis minutiflora, in a Venezuelan savanna.
Plant Ecol. 167, 31–43.
Baruch, Z., 1996. Ecophysiological aspects of the invasion by
African grasses and their impact on biodiversity and
function of Neotropical savannas. In: Solbrig, O.T.,
Medina, E., Silva, J.F. (Eds.), Biodiversity and Savanna
Ecosystem Processes: A global Perspective, Ecological
Studies, vol. 121. Springer, Berlin, pp. 79–93.
Baruch, Z., Fernández, D.S., 1993. Water relations of native
and introduced C4 grasses in a neotropical savanna.
Oecologia 96, 179–185.
Baruch, Z., Gómez, J.A., 1996. Dynamics of nutrient and
energy concentration and construction cost in a native and
two alien C4 grasses from two Neotropical savannas. Plant
Soil 181, 175–184.
Beard, J.S., 1953. The savanna vegetation of Northern South
America. Ecol. Monogr. 23, 149–215.
Berry, P.E., Holst, B.K., Yatskievych, K. (Eds.), 1995. Flora
of the Venezuelan Guayana. Vol. I. Introduction. Missouri
Botanical Garden, St. Louis and Timber Press, Portland,
OR, pp. 97–160.
(Estudio ecológico de la region alrededor de Calabozo, Edo.
Guárico). Bol. Soc. Venez. Cien. Nat. 23 (102), 139–206.
Cole, M.M., 1986. The Savannas: Biogeography and Geobo-
tany. Academic Press, London.
Filgueiras, T.S., 2002. Herbaceous plant communities. In:
Oliveira, P.S., Marquis, R.J. (Eds.), The Cerrados of Brazil.
Columbia University Press, New York, pp. 121–139.
Frost, P., Medina, E., Menaut, J.C., Solbrig, O.T., Swift, M.,
Walker, B., 1986. Responses of savannas to stress and
disturbance. Biol. Int. (Special Issue 10. International
Union of Biological Sciences, Paris).
Goldsmith, F.B., 1974. Multivariate analyses of tropical
grassland communities in Mato Grosso, Brazil. J. Biogeogr.
1, 111–122.
Goodland, R.J., 1966. South American savannas: comparative
studies Llanos & Guyana. McGill University Savanna
Research Project Series 5, Montreal, Canada, pp. 1–52.
Goodland, R., Pollard, R., 1973. The Brazilian cerrado
vegetation: a fertility gradient. J. Ecol. 61, 219–224.
Hill, M.O., 1979. TWINSPAN: A FORTRAN program for
arranging multivariate data in an ordered two-way table by
classification of the individuals and attributes. Cornell
Ecology Programs Series. Cornell University, Ithaca, NY.
Hoffmann, W.A., Jackson, R.B., 2000. Vegetation climate
feedbacks in the conversion of tropical savanna to grass-
land. J. Climate (Am. Met. Soc.) 13, 1593–1602.
 ARTICLE IN PRESS
64 Z. Baruch / Flora 200 (2005) 49–64
Hoffmann, W.A., Schroeder, W., Jackson, R.B., 2002. Positive
feedbacks of fire, climate and vegetation and the conversion
of tropical savanna. Geophys. Res. Lett. 29 (22), 2052.
Huber, O., 1987. Neotropical savannas: their flora and
vegetation. Trends Ecol. Evol. 2, 67–71.
Huber, O., 1995. Vegetation. In: Berry, P.E., Holst, B.K.,
Yatskievych, K. (Eds.), Flora of the Venezuelan Guayana.
Vol. I. Introduction. Missouri Botanical Garden, St. Louis
and Timber Press, Portland, OR, pp. 97–160.
Jackson, M., 1982. Análisis Quı́mico de Suelos. Prentice-Hall,
Englewood Cliffs. New York.
Jongman, R.H., ter Braak, C.J.F., van Tongeren, O.F.R.,
1987. Data Analysis in Community and Landscape
Ecology. Pudoc, Wageningen.
Klink, C.A., Moreira, A.G., Solbrig, O.T., 1993. Ecological
impact of agricultural development in the Brazilian
cerrados. In: Young, M.D., Solbrig, O.T. (Eds.), The
World’s Savannas. UNESCO. The Parthenon Publishing
Group, Paris, pp. 259–282.
Lopes, A.S., Cox, F.R., 1977. Cerrado vegetation in Brazil: an
edaphic gradient. Agron. J. 69, 28–31.
McCune, B., Mefford, M.J., 1999. PC-ORD. Multivariate
Analysis of Ecological data, Version 4. MjM Software
Design, Gleneden Beach, Oregon, USA.
Medina, E., Silva, J., 1990. Savannas of northern South
America: a steady regulated by water-fire interactions on a
background of low nutrient availability. J. Biogeogr. 17,
403–413.
Melillo, J.M., McGuire, A.D., Kocklighter, D.W., Moore, B.,
Vorosmarty, C., Scloss, A.L., 1993. Global climate change
and terrestrial net primary production. Nature 363,
234–240.
Miranda, I.S., Absy, M.L., Rebelo, G.H., 2003. Community
structure of woody plants of Roraima Savannahs, Brazil.
Plant Ecol. 164, 109–123.
Moreira, A.G., 2000. Effects of fire protection on savanna
structure in Central Brazil. J. Biogeogr. 27, 1021–1029.
Mueller-Dombois, D., Ellenberg, H., 1974. Aims and Methods
of Vegetation Ecology. Wiley, New York.
Murphy, J., Riley, J.P., 1963. A modified simple method for
the determination of phosphate in natural waters. Anal.
Chem. Acta 27, 31–35.
Oliveira-Filho, A.T., Ratter, J.A., 2002. Vegetation physiog-
nomies and woody flora of the cerrado biome. In: Oliveira,
P.S., Marquis, R.J. (Eds.), The Cerrados of Brazil.
Columbia University Press, New York, pp. 91–120.
Parsons, J.J., 1972. Spread of African pasture grasses to the
American tropics. J. Range Manage. 25, 12–17.
Pieters, A., Baruch, Z., 1997. Effect of soil depth and fertility
on biomass and nutrient allocation in Jaraguagrass.
J. Range Manage. 50, 268–273.
Pivello, V.R., Shida, C.N., Meirelles, S.T., 1999. Alien grasses
in Brazilian savannas: a threat to the biodiversity.
Biodivers. Conserv. 8, 1281–1294.
Ramia, M., 1967. Tipos de sabanas en los llanos de Venezuela.
Bol. Soc. Venez. Cien. Nat. 27 (112), 264–288.
Ramia, M., 1974. Plantas de las Sabanas Llaneras. Monte
Avila Editores, Caracas.
Ramia, M., 1993. Ecologı́a de las Sabanas del Estado
Cojedes: Relaciones Suelo-Vegetación en Sabanas Secas.
CONICIT-Fundacion La Salle de Ciencias Naturales,
Caracas.
San José, J.J., Fariñas, M.R., 1991. Temporal changes in the
structure of a Trachypogon savanna protected for 25 years.
Int. J. Ecol. 12, 237–247.
San José, J.J., Montes, R., Mazorra, M., 1998. The nature of
savanna heterogeneity in the Orinoco Basin. Global Ecol.
Biogeogr. 7, 441–451.
Sarmiento, G., 1983. The savannas of tropical America.
In: Bourliere, F. (Ed.), Tropical Savannas. Eco-
systems of the World, vol. 13. Elsevier, Amsterdam,
pp. 245–288.
Sarmiento, G., Monasterio, M., 1969. Studies on the savanna
vegetation of the Venezuelan llanos. I. The use of
association analysis. J. Ecol. 57, 579–598.
Saugier, B., Roy, J., Mooney, H.A., 2001. Estimations of
global terrestrial productivity: converging toward a single
number? In: Roy, J., Saugier, B., Mooney, H.A. (Eds.),
Terrestrial Global Productivity. Academic Press, San
Diego, pp. 543–557.
Silva, J.F., 1996. Biodiversity and stability in tropical
savannas. In: Solbrig, O.T., Medina, E., Silva, J.F. (Eds.),
Biodiversity and Savanna Ecosystem Processes: A global
perspective. Ecological Studies, vol. 121. Springer, Berlin,
pp. 161–174.
Steyermark, J.A., Huber, O., 1978. Flora del Avila. Sociedad
Venezolana de Ciencias Naturales Caracas.
Susach, F., 1989. Caracterización y clasificación fitosociológi-
ca de la vegetación de sabanas del sector oriental de los
Llanos Centrales Bajos Venezolanos. Acta Biol. Venez. 12,
1–54.
Tamayo, F., 1943. Notas de ecologı́a venezolana. Proceso de
despoblación y reposición vegetal de las colinas de Caracas.
Instituto Pedagógico, Caracas.
ter Braak, C.J.F., 1987. The analysis of vegetation–environ-
ment relationships by canonical correspondence analysis.
Vegetatio 69, 69–77.
Thomas, D., Vera, R.R., Lascano, C., Fisher, M.J., 1990.
Use and improvement of pastures in neotropical
savannas. In: Sarmiento, G. (Ed.), Las Sabanas Amer-
icanas: Aspectos de su Biogeografı́a, Ecologı́a y Utilización.
Fondo Editorial Acta Cientı́fica Venezolana, Caracas,
pp. 141–162.
van Donselaar, J., 1965. An ecological and phyto-
geographic study of northern Surinam savannas. Wentia
14, 1–163.
Vareschi, V., 1970. Las sabanas del Valle de Caracas. Acta
Bot. Venez. 4, 427–522.
Vareschi, V., 1992. Ecologı́a de la Vegetación Tropical.
Sociedad Venezolana de Ciencias Naturales, Caracas.
Velásquez, J., 1965. Estudio fitosociológico acerca de los
pastizales de las sabanas de Calabozo, Edo. Guárico. Bol.
Soc. Venez. Cien. Nat. 26 (109), 59–101.
Walter, H., 1973. Vegetation of the Earth. Springer, New
York.
Williams, D.G., Baruch, Z., 2000. African grass invasion in the
Americas: ecosystem consequences and the role of ecophy-
siology. Biol. Invas. 2, 123–140.