The Vegetation of Falcn State, Venezuela

Author(s): Silvia Matteucci

Source: Vegetatio, Vol. 70, No. 2 (Jun. 15, 1987), pp. 67-91
Published by: Springer
Stable URL: http://www.jstor.org/stable/20038134
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1987
70: 67-91,
Dordrecht
Junk Publishers,

Vegetatio
? Dr W.

The

of Falcon

vegetation

in the Netherlands

Printed

67

Venezuela*

State,

Silvia Matteucci**
de Producci?n
Vegetal, Universidad
Departamento
tado 7434, Coro, Falc?n,
Venezuela 4101

Formation-series,

Classification,

Keywords:

Nacional

Experimental

Dominance-type,

Francisco

Physiognomy,

de Miranda,

Apar

Venezuela

Abstract
first approximation
A semi-detailed
study of the vegetation of an heterogeneous
area, located in the Caribbe
an region is presented. Field information on vegetation
structure and function, flora and environmental
fac
tors was collected in a systematic way in order to reduce subjective decisions as much as possible.
Indepen
were developed
and floristic classifications
through the analysis of field data. The
physiognomic
a
to
classification
is
understand
wide
artificial, easy
physiognomic
by
variety of users. It facilitates the iden
tification and description of land units, as well as the monitoring
of vegetation changes. The floristic classifi
was obtained from tabular comparison.
The relationship between both
cation, based on dominance-types,
dent

is analyzed. In order to disclose the temporal and spacial relationships among vegetation units,
for extraregional
the results were translated into Beard's formation
series. The
comparisons,
between the physiognomyand dominance-types
and the formations are analyzed. The vegetation

classifications

and to allow
relations

in terms of Beard's

is described
are discussed.

formations.

The relationships
to those of previous

results are compared

The

Introduction
the vegetation
Although
bean region has been
Robbins,

1953;

Stoffers,
Caribbean

1956),

of the parts of the Carib

described
(e.g., Asprey &

Beard,
little has

1949;
been

Howard,

1973;
reported on the
of northern Venezuela.
This

vegetation

between

Nomenclature

able on request).

Herbarium

National
um

(MY)

and

** I am
deeply
actively

Dr D. Billings
editorial
work
Council

was

in the Central

(VEN),

in the reference

de Coro,

nol?gico

pated

et al. (1979a,
of species follows Matteucci
have been deposited
Voucher
specimens
collection

University
of

the

avail
in the

Herbari

Instituto

Tec

to A. Colma

and L. Pia, who partici

at all stages. I am very grateful
to
on the manuscript,
for his comments
and to the
in the project

secretariat
supported
(CONICIT).

for helpful
by

and environment

land use
regions
M.A.C.,
Tamayo,

purposes
(O.T.E.E.,
1958) or in larger
that include Falcon State (Hueck,
1960;
1961; Pittier,
1920; Smith et al,
1973;

are of a
these accounts
1958). However,
nature and the vegetation
are
types
only
described. Most of the studies rely on aeri

vaguely
al photographs
with very little field checking. Fre
map of Venezuela
quently, the biogeographical
by
Ewel & Madriz
is
used
for
the
characteriza
(1968)
tion of the local vegetation. However,
this is a cli
matic map and the communities
described within

Falc?n.

indebted

pattern

paper is the first attempt to produce a comprehen

sive semi-detailed
study of the vegetation of Falcon
State. Other
studies have been done for specific

general
*

vegetation

studies.

improvements

the Venezuelan

of

the text. Field

National

Research

the life zones of Falcon

the actual
most

nor with

State coincide
the potential

neither with
vegetation,

in

cases.

The present account

is based on a systematic
field study of the vegetation and flora as part of a

68
regional survey (Matteucci et ai, 1985) which was
for research
to provide
information
the baseline
and development
planning.
and floristic classifications

Simple physiognomic
were developed,
that

by non-specialists.
easily understood
of informa
the systematic collection
of land
and definition
tion and the identification

would

be

These

allow

units.

These

classifications

were obtained

the results
for extraregional
comparisons,
in terms of Beard's formation-series

were analyzed

system (Beard, 1944, 1955). In dry climates, differ

ences in topography,
exposure and soils, as well as
rainfall seasonality have greater influence on vege
tation

than mean

pattern

Beard's

takes

approach

annual

into account

precipitation.
this fact in

to environmental
features. It
relating physiognomy
re
also provides a means for detecting successional
to
situations.
local
it
is
and
lations,
applicable
The natural vegetation has almost entirely disap
peared in Falcon. It is likely that most of the pres
ent

closed

forests

and woodlands
succession.

of

late
represent
stable
Different

stages
secondary
communities may have replaced the natural vegeta
tion types, due to the fact that changes in habitat
recovery in a tropical
may be faster than vegetation

and climatic conditions

the topographic
Even
in
Falcon.
though it is likely that
prevailing
not
exist here, it is possible
do
climax communities
to detect apparently stable, integrated, mature com

area with

munities, which
climax (Beard,

of
comply with Beard's definition
eco
of
lack
the
However
1944).
of

and

local

species
knowledge
physiological
renders
facts about land occupation
historical
difficult
Steenis,
1961).
(Van
interpretation
This paper also aims at presenting a method

of
the
to

of an extensive
study and interpret the vegetation
region at a first approximation.
heterogeneous

The

Valleys.
Valleys and Eastern Maritime
and soils
of geology,
geomorphology
elsewhere
et ai,
have been published
(Matteucci
&
In
Colma
Creta
the
1979b;
Matteucci,
1982).

Central

Details

the area was occupied by a g?osynclinal

covered
direction,
stretching in a ENE-WSW
sea. Several episodes of continental
continental
ceous,

independent
a description

in order to provide
ly and combined
of the present vegetation
(Matteucci et al., 1979a).
in order to obtain a temporal and spatial
However,
and
of the community
distribution,
interpretation
to allow

prises 24750 sq. km of land with extreme varia

and history of human
tions in relief, topography
are
five
There
provinces:
physiographic
occupation.
Falcon Ranges,
Coastal Plains, Coastal Piedmont,

interbedded shale and

produced
deposition
and lime
sandstone, with layers of conglomerate
stone. Orogenetic
activity started in the Lower Mio

marine

the uplifting and tilting of the Coastal

and
ended in the Pliocene. The deposi
Piedmont,
are
overlain by Quaternary
tional plains
deposits,
cene, with

with

The

State

borders

the Caribbean

Sea.

It com

exposed

on

the erosional

the Paraguan?
Plains
comprise
and
coastal
the
isthmus
the
fringe to the

Coastal

Pen?nsula,
west. The

is low, with extensive floodplains,

of the mainland,
central portion
is raised and denuded, with exposed stony
is
surfaces. The highest altitude
(815 m altitude)
The
found on an isolated hill in the Peninsula.

except
which

relief

in the

consists of
formed by recent sand deposits,
areas
rivers are
The
dunes.
of
with
beaches
sandy
with
flashfloods
and
carry
intermittent,
suspended
in the rainy season. In most of the prov
sediments
to fine-textured,
of the
ince the soils are mediumshore,

and Orthents
(Calcic
Argids,
on
soils
The
and
Yermosols,
Lithosols).
Xerosols,
the coastal fringe and isthmus belong to the subor
suborders

Orthids,

der Psamments
The Coastal

(Eutric Regosols).
Piedmont
constitutes

a belt of tran

and
erosional
relief, with
topographic
Coast
the
between
located
landforms,
depositional
It varies in eleva
al Plains and the Falcon Ranges.
sitional

tion from

100 to 400 m towards

and consists

its southern border,

and dis
of hogbacks
soils are shallow and lithic

of a succession

low ridges. The

except in the few valleys and depres
(Lithosols),
sions, where the suborder Orthids (Haplic and Luv

sected

The
Falcon

sediments

Tertiary

landforms.

ic Xerosols)

study area

belt
by a
and

Falcon

predominates.
Ranges Province

the State and comprises

occupies
three ENE-WSW

44 ?/o of
oriented

69

parallel
northern

ridges separated
by wide
valleys. The
are
and middle
divided
into three
ridges

The

coastal plains,
show a climatic

main mountain-masses

winds,
such a maritime

which

would

by gorges and valleys along

some rivers flow northwards. The slopes are

the relief is drastic, with

and v-shaped ravines. The alti
tude ranges from 200 to 1500 m. To the east,
remnants are found within
mountain
the Eastern
steep

(25-50 %)
divides
knife-edge

and

Valleys. On the top of these, as well as on

the easterly mountain-mass
of the northerly ridge,
karst landscapes
formed by calcareous masses
of
m
of exposed limestone have developed.
up to 350
The soils are heterogeneous:
and
mostly Orthids

Maritime

on the
and Lithosols)
(Calcic Xerosols
thick
Rendolls
slopes;
organic
(Rendzinas) on the
mountain
summits, and Tropepts and Orthids
(Eu
tric Cambisols
and Haplic Yermosols)
in the val
Orthents

leys.

The

Central

the
encompasses
Valleys Province
the mountain
syncline
valleys between
ranges.
relief is low and mostly
flat or un
Topographic
dulating, except for some low foothills
ed, narrow buttes. The soils, shallow

and elongat
and covered

by gravel, belong to the suborder Orthents

(Calcic
Xerosols
and Lithosols).
In the valleys and dried
the soils are fine-textured
and
up water courses,

to surface

subjected

At

anomaly.
location, abundant

trade

this latitude,

in

precipitation
be expected. However,
limited and erratic
rainfall prevails. The possible cause of this anomaly
is the stress that arises from the interaction between
and the cool Caribbean
topography
Sea, which
forces the air masses to diverge and subside over the

dry area

(Lahey, 1973). On the mountain

ranges
rainfall occurs with rainshadows on the
orographie
lee sides. The summits are subjected
to frequent
cloud cover. Rain is also less sparse in those valleys

which

to

open

the

comes

Rainfall

east-north-east.

as heavy

tion. Rainfall
detailed

showers

of short dura

characterizes

seasonality
analysis of local climatic

the area. A

data has shown

that

the ratio of potential

to
?vapotranspiration
3.56
is
&
temperature
Colma,
(Matteucci
1986).
of the
Thus, in order to give a realistic description
in such a
climate, the diagrams were constructed
to 35.6 mm
way that 10 ?C are made to correspond
precipitation
(Fig. 1). On the coastal plains and
central valleys mean
annual precipitation
ranges
from 142 to 492 mm and is concentrated
in a short

overlain

salt deposits, of the subord

by gypsiferous
ers Fluvent
and Orthids
and Gypsic
(Fluvisols

,A6UA CLARA?85)

(25)
28,7?C 463mm. .TUCACAS

Xerosols).
The Eastern Maritime
sins of the four main

the ba
Valleys comprise
rivers that flow to the east and
are bor

dered

from the Falcon Ranges.

by low divides formed

by undulating

plains

or mountain

The

north

300 m

remnants.

The basins
altitude

high
ranges

to the foothills
to 0 m at the
shoreline. Slopes are gentle and near the coast the
flood
plains become
temporarily or permanently
from

ed. Soils

next

JT

' ' ' ' ' ' '

'M'A rM J J A S 0 N D

1ARAURIMA(35m)
19

" '
'
J F'M' A" M"J J A'S' O'N" D

27,6?C 1527mm SOCOPO(520)

16

24,5?C 1Il76mmi

form a mosaic

of patches of Usterts and

Argids (Chromic Vertisols and Luvic Yermosols) on
the hills and undulating
plains, with fine-textured
and
saline
clayey
patches of Orthids and Aquents
on the
and Eutric Gleysols)
in the southern portion,
valleys, whereas
and Aquents
Tropepts, Fluvents,
(Eutric Fluvisols
and Gleysols) occupy the river margins, and Ustalfs
(Haplic

Yermosols

northerly

and Usterts

(Dystic Nitosols

predominate

elsewhere.

and Ferric Luvisols)

J7'MTA^M'

J'J

'
A'S

'O'N'D

'
J' F' M'A'M" J J'A'S

0 KTD

1. Climate-diagrams,
with
in ?C on
temperature
axis and precipitation
in mm on the right one.
see text.
planation

Fig.

the

left

vertical

For

ex

70
rainy season (Fig. la). The high ?vapotranspiration
rate produces drought all year round. In the Mari
time Valleys, the rainy season is longer (two to four
mean annual precipitation
ranges from
months);
600 to 1700 mm

Fosberg's
recorded

and there are two to nine humid

The

is a N-S

There

mean

annual

precipitation

mic

coefficient

with

lianas,

Leaf

Data

study

tation.
ed

depicts
form was

area was

Land

on

stratified

segments

(0.25

through deductive
photointerpre
sq. km minimal
area) were delineat

1:60000

panchromatic

scale

units. After

556 photo-interpretation

field

on a representative
each unit. In the field, the
site within
to confirm
segment was traversed as far as roads permitted
and

site representativeness
spection of a sample
data

check

up

stand of around

as well

as

information

0.5 ha, physiognomic

on the environment

gin.
The

land

Table

on a check

1. Plant

Closed:

crowns

of

Open:

level was

identified
The

fall

of

shoots

or shoots

their diameter
sparse:

Data

Through

the relative

for each

shows

the

is represented
cover degree.

interval

of

the

of

abundance

stand

(Fig. 2).

by the horizontal
The

line

vertical
Each

leafy portion.
growth
to facilitate visual
by a colour and a symbol,
form was ob
percent cover for each growth

the summation

check

sheet

information

included

hydrology

and

land

use

on

soils,

(Matteucci

relief,
et al.,

analysis
visual

comparison,

the profile

diagrams

were

grouped

Species
Continuous

touch

>75%

(6):

do not

touch,

but

Scattered

dominates

Sporadic

(5): 50-75%

15-25%

Rare (3):

Very
the

Interrupted

25-50%

(4):

apart
the substrate

landscape

Thorniness

is, by

synusia

This

species.

is syn
species
was deter

deciduous

species.
was constructed

that

from

determined

and evergreen

intervals.

they cover at least 30% of the surface

than twice
crowns or shoots more
Sparse:

Very

of the same elements

groupings

or overlap
crowns

literature.

of a
by means
use of two differ

and
were

form

Growth

and
obser

repeated

1985).

sheet. Random

cover

cacti,

climbers,

in

have been
sampling would
in such an extensive and
and time consuming
extremely difficult
heterogeneous
region and was not attempted.
recorded

length

microtopography,

the

field

limits. After

the

1). The

forbs,

of the mean cover degree of the spe

in
synusia with less than 3 % cover were not included
the profile, but were listed as present or absent on the right mar

crop

reconnaissance,

shrubs,
column

cies. The

in

resulting

photographs,

(Table

the dry season.

of each

the height

comparisons.
tained from

units
and 213 vegetation
land units were discarded
and pasture
on the aerial photographs
were recognized.
Plots were marked

floristic

leaf
of

same way;

height

line, whose

collection

The

because

thorny vs non-thorny
A profile
diagram
The mean

estimated

of deciduous

at the onset
in the

than 5 m),

was

at the community

abundance

sense

species
were: tall

the estimations.

periodicity

chronized

each

considered

herbaceous
succulents,
was obtained
from
persistence
or from the local
the seasons,

to check

makes

Methods

were

shape

of

stage

small

scale
Braun-Blanquet
to assess two different

the relative

mined

to

according

and

hemi-parasites,

epiphytes,

cover-abundance

species

ent scales
served

layers,
the

of

were col
specimens
not possible. Growth

listed. Voucher

low trees (shorter

succulents,

modified

an altither

size,

and phenological
leaf persistence
The growth form categories

tuft plants. Leaf

vations
through
The

up the mountains
of 0.57.

texture

recorded.

shrubby

high temperatures
prevail
the year, though day-night temperature
throughout
differences may reach up to 14?C. On the Coastal
is 28 ?C to 29 ?C,
Plains mean annual temperature

as its spacing

in the field was

identification

graminoids,

(Fig. Id).
uniform
Remarkably

and it decreases

(Table

trees (5 m or taller),

1250 mm

1). Leaf

synusia.
present were

species

form,
were

as well

synusia,

of

height

for each

lected when

to

750

from

ranges

of each
scale

number

and minimum

maximum

each,

were

considered

features

of

height

leafy portion

humidity gradient, and the

northern
(Fig. lb) than the
fringe is less humid
On
the Mountain
southern
portion
(Fig. lc).
and
the
southern
extremity of the western
Ranges
Coastal
Plains
there are two rainy seasons and

months.

structural

The
mean

(1):

1-

rare

5-15%

(2):

5%
Present

+ ):

<

1%

71
CENSO 45
MITARE

9H

in common.
The dominance-types
the genus Eugenia
or the most
after the dominant
frequent
species

and have
were

BDSE

in

named

the cluster.

8
7

1 6

Data

Profile

UJ 4i

Beard's

s=0
g=+
H =+
E=+
P=0
En=0

H
2
I
"I?r?r?i-1-1-1-1-1-1
10 20 30 40
COVER

interpretation
diagrams
(1944,

to serai stages and

with Beard's
correspond
descriptions
to which
to the formations
and associations
they
assigned
appear to be related, on the basis of floristic and physiognomic

meet

were

environmental
elements,
the concept of potential
50

60

70

DEGREE

80

90

100

Ellenberg,
The

(%)

2. Profile
A = trees; a = shrubs;
SC = column
diagram;
=
=
=
L
sa
s
h = forbs;
cacti;
lianas;
succulents;
shrubby cacti;
=
H = hemi-parasites;
P = tuft
g
grasses;
plants;
=
En = herbaceous
BDSE = closed ever
climbers; E
epiphytes;

Fig.

life form

structure
in clusters
and
these were
of similar
vegetation
to the community
function
according
segregated
(periodicity
to produce a physiognomic
and thorniness)
hierarchical
classifi
from the
cation. The definitions
of the classes were formulated
shared

by the members

to facilitate

graphs
An

of each

future vegetation

was

classification

or by field
independent

by

checking.
floristic

classification

with

comparison

pre

in the field. The

typification

validated

key was

cluster. A

based

land photo
on

dominant

from the species

lists.
1978) was developed
After each field trip, these were added to a raw table. The domi
nant and co-dominant
and the columns
species were underlined
species

(Whittaker,

were

to group together all the relev?s that contained

arranged
the same dominant
group (Whit
species or the same commodal
dominant
when
its abso
taker, 1978). A species was considered
was

lute coverage

16 % or over and

considered

species
age did not differ in more
and rows were alternatively
leve clusters

In some

composition.

either

(no species
it is very

dense,

floristic

times

to show

applying

(Mueller-Dombois

&

were

spectra for the formations

cover of each growth
were

features

taken

calculated

from

form.

into account

to assign

the

to the formations.

and have

is sparse

similar

to. determine
and diverse

16 % of the sample plot) or because

co
rich and diverse
(there are too many
to the former
relev?s conforming
situation
than

affinity,
and

given

the minimum

by

with

which

they had

the

the maximum

number

of

of single

Classification

The physiognomic
The

systems
classification

first order of classification

the pri
comprises
the
domi
characterized by

structural groups,
the vertical
growth form, which determines
are segregated
structure. These
into structural
to the canopy
groups (the second level) according
cover.
to the
The third level corresponds
percent

mary
nant

physiognomy-type,
the community
structural

which
function

47 physiognomy-types
Falcon State.
Forests

includes consideration
(Fig. 3). Five

18 structural

groups,

primary

groups

have been

of

distinguished

and

in

and woodlands

the communi
comprise
in the
by trees (Fig. 4); however,
there is at least one tree layer > 5 m high,

ties dominated
forests

re

species. The relev?s

were grouped
to the rich dense communities
to
to the most
humid environments
they correspond

species

corresponding
since
gether,

was over

from each other. Columns

the vegetation

The
dominants).
were assigned
to the dominance-type
greatest
common

coverage

it is not possible

situations

because

cover more

than 30 %

several
rearranged
both share dominants

that would

dominance,

its relative

the accompanying
species. Two or more
co-dominant
when
their relative cover

30 % of that of any of
were

and

geography,

percent

Environmental
stands

Results:

pared

and

vegetation

forest.

thorny

features

factors
natural

1974).

the relative

green

were compared
to profile diagrams
based on
Those communities
that do not
1955) definitions.

and

one

or

two

lower

layers.

In

the

woodlands,

is <5 m

and vegetation
height of trees
the tree species
layering is obscured.
Frequently
have a shrubby appearance.
Shrublands
include
average

degree of cover by shrubs ex

ceeds that of trees by at least 30%. The 'cardonales'
in which the column
comprise those communities
cacti occupy at least 30 % of the canopy on a plant
communities

cover

basis.

those whose

in which

Closed
dominant

communities

are defined

as

synusia covers at least 75 %

72
ment

to

The
are

the more

stable

forests, which

scattered

community-types.

35.5 % of

occupy

two-

The

everywhere.

the State,
and

three

layered evergreen closed forests are located at an al

titude of 200 to 800 m on the hills near the coast
line, up to 1 500 m on the Falcon Ranges Province,
and at 20 to 100 m in the southern extremity of the
Valleys. They are associated with deep or
or
soils and high mean annual precipitation
cover. The unilayered
forests located on the

Maritime
ganic
cloud

lower slopes of the mountain

ranges and to the
south of the Western Coastal Plains are deciduous,
open or closed and rather low (5 to 10m). Those
at the eastern extremity of the mountain
ranges and
in the Maritime Valleys, where climate is subhumid,
or
(8 to 15m) and denser, deciduous
es
with
noticeable
trees,
evergreen
semideciduous,

are higher

season.

pecially during the dry

semi-arid Coastal Plains,
to the river banks.

In the arid

and

are confined

the forests

The woodlands
(18.5% of the State) are concen
trated in the semi-arid zone, within all the Physio
lies within
graphic Provinces. The largest extension
in the Paraguan?
Plains,
specially
%
7.1
subhumid
of
the
region is
Only

the Coastal
Pen?nsula.
3. Physiognomie
classification
system. The physiognomy
as follows:
are
the first letter
letters
types
by capital
depicted
to the vertical structure: B = forest; M = woodland;
corresponds
A = shrubland;
VH = herbaceous
C = cardonal;
vegetation.
structure:
to the horizontal
letter corresponds
The
second
third capital
letter
R = open; Dt = desertic.
The
D = closed;
S = evergreen; D = deciduous;
refers to the foliage periodicity:
of
the abundance
Sd = mixed.
letter represents
The
fourth
=
=
The superscripts
non-thorny.
thorny; I
thorny structures: E
2 & 3 in the forests refer to the number of tree strata, the absence

Fig.

indicates
that there is only one tree layer; the low
of superscript
er case letters following
the superscript
refers to the height of the
=
a = canopy
lower
than 5m;b
forest canopy:
canopy
higher
than 5 m.

of the surface; in open communities

plant cover of
the dominant
synusia ranges from 75 to 50 % while
in desert vegetation
plant cover is less than 50 %
and the substratum

dominates

the landscape.
(34 %) are represent

Sixteen physiognomy-types
unit each (Fig. 4);
ed in only a single physiognomic
to
degraded impoverished commu
they correspond
nities.

In most

physiognomic

the presence of floristic or

relict elements allows their assign
cases,

most

occupied by woodlands;
to disturbed
forests.
The

shrublands

most

The

occupy

of which

only 7.6 % of the State.

are

communities

extensive

Pen?nsula

and

correspond

located

in the

in the Maritime

Valleys.
Paraguan?
The former are open or desertic xerophytic commu
nities dominated
by evergreen thorny species. The
zone shrublands, which are deciduous
subhumid
and
and

occupy

non-thorny,
represent

serai

stages

abandoned
in

pastureland

secondary

succes

sion.

'cardonales' (3 % of the State) occur in the

are
zone
only. The most extensive communities
dry
Pen?nsula
located in the north of the Paraguan?
and on the western alluvial plains; there are small
The

Most
of
the Central Depressions.
patches within
them develop on flat, well drained soils.
The herbaceous
occupies a very
community-type
reduced area (0.24 % of the State) and is scattered
in patches on the coastal fringe. It is dominated
by
forbs with very few grasses, thus they cannot be in
cluded in the savannas.
Around

24 % of

the study area

is occupied

by

73

|^75%
Fig.

4. Growth

growth
(each

form

of which

may

ing with

fire.

of

correspond

either

pastureland,

of

spectra

form. The meaning

These

Pf

50-75?/

i 25-50%

which

fs*

:l%

00%

The symbols
the cover degree of each
each of the physiognomy-types.
represent
row
3.
in
last
of physiognomic
units
is
indicate
the
number
in
The
the
symbols
Fig.
figures
given
to one or more photo-interpretation
units) in each physiognomy-type.

stands

representing

sown or developed
are man-made

forest

11-5%

letter

after clear

savannas,

with

of grasses,
predominance
grazed by cattle and
maintained
by periodical
burning. To the east, in
the Maritime Valleys, the savanna constitutes a ma
trix within

[115-25% Kl 5-15%

remnants

are found.

in this dominance-type
and 100 were present
On the other
and related dominance-types.

in this

hand,
from each of the
only one species was discarded
Caste la erecta, Prosopis juliflora-Ritterocereus
and
Ritterocereus
spp dominance-types.
is some overlap in the woody species pres
ence and there are groups of dominance-types
that
There

differ more
The floristic

classification

The

classification

floristic

with respect to the relative abundance

of species than to their presence. This is noticeable
that is,
the most xeric dominance-types;
among
includes

15 clusters

and

from the Ritterocereus

spp dominance-type

to the

The clusters represent the dominance

types, arranged in a gradient from the humid to the
xeric types in Table 2, which shows the woody spe
cies composition
of each type. The figures cor
to
the presence percentage of the species in
respond

corlarla
right in the table. For example, Caesalpinia
in these four
has a high presence
percentage
clusters, but is dominant
only in the C. corlarla

the group. The table does not include all the species
recorded, since those present in one or two to three
related dominance-types
less than 50 %
holding

abundant.

9 outliers.

presence in each, were discarded. Thus, of almost

600 woody species recorded, only 144 are listed. In
the Eugenia
416 woody
spp dominance-type,
spe
cies were recorded, of which 212 were present only

Acacia
dominance-type;
four types, with varying

tortuosa

is present in the
but it is never
percentages,

is exclusive to any of the

12
Only
species are not found in
dominance-types.
the humid types (from the Apoplanesia
cryptopeta
to the left in the table) and
la dominance-type
106 species plus those discarded, are not present in
None

the xeric

of

the species

types.

Six

species

are present

in the

74
in each
the presence
The figures depict
for each species
of the dominance-types.
2. Woody
percentage
species composition
as a percentage
on the basis of the number of stands where
the species occurred
of the number of stands
determined
dominance-type,
=
are labeled: Pj = Prosopis
The dominance-types
Ce = Castela
cori
in the respective dominance-type.
erecta', Ce
juliflora;
Caesalpinia
=
=
=
=
=
RR
Ritterocereus
Cercidium
Ch
Crot?n
Tabebuia
spp; AA
spp; P
praecox;
heliaster; Cp
Aspidosperma
aria; Tb
billbergii;
= Bourreria
EE = Eugenia
brasiliensis-Bulnesia
R = P. juliflora-Ritterocereus
spp;
arb?rea',
spp; Be
cumanensis-Phyllostylon
=
Pd = Pithecellobium
Ac = Apoplanesia
hastata; Zp
pterota-Machaerium
Zanthoxyllum
spp-Eugenia
dulce-Capparis
cryptopetala;
=
spp.
spp; PP
Pilocarpus
Table

EE

TYPE
DOMINANCE

SPECIES

Zp

PP

Ac

Ch

Pd

Haematoxylon brasiletto
Castela
Cercidium

erecta
praecox

Lippia origanoides
2 izyphus saeri
Lye ium nodosum
Guaiacum officinale
Capparis I inear is

14
29
29
43
14

arenosa
Mimosa
Geoffraea
spinosa

Cassia biflora
Pithecellobium
platylobum
Diphysa carthagenensis

Pereskia
Bursera

guamacho
tomentosa

Cissus tri fol iata

Manihot carthaginensis
Bur sera

karste?iana
caracasana

Jacquinia

Ruprecht ia ramiflora

Acacia

tortuosa

Caesalpinia cor iaria

Ritterocereus
spp
Prosopis julif lora
odorat issima
Capparis
Pitnecellobium
dulce
Cissus sieyoides
Crot?n heliaster
Tabebuia billberqf

10
13
11
7
33

Bulnesia

arb?rea"
mor
Cephalocereus
hastata
Capparis

Malpighia

itzi

anus

spp

cumanensis
Bourreria
Cassia
emarginata

Bumeli a obt?si fol ia

mo!1 is
Caesalpinia
Rand ia gaumeri
tr?mula
Casearia
Astron?um
graveolens
revoluta
Jacquinia

Tal isi a oliuaeformis

Aspidosperma

cuspa

Guapira spp
Pilocarpus spp
brasiliensis
Phyllostylon
Capparis f?exuosa
Zanthoxylum pterota
Phyllanthus botryanthus
Cordia

curassavica

Machaon ia ottonis
Gyrocarpus
Pseudobombax

americanus
septenatum

Plumer ia oudica
Capparis tenu isil iqua
Acacia tamarindi folia
Zanthoxylum monophyllum
Mor ison ia americana
corail
ina
Arrabidaea
frutescens
Myrospermum

Celtis

iguanea

Matelea
Chioccoca

mari tima
alba

Cestrum al tern ifolium

Manihot aff. brachyloba
Tabebuia

chrysea

20
13
13
2?
27
7
7
13
?
20
7
40
13
13
13
73
7
7
13
40
7
7
13
33
20
33
7
13
40
13
7
13
7
33
7

43
14
14
2?
14
2?
71
57
57
14
29
57
5714
29
71
57

67
33
33
33
33
33

20
40
40
20
20
20

67
67
67
67
67
67
100
100
100
67
67
33
33
67

40
20
40
40
80

60
80
60
80
20
60
20

33
33

40

33

40

29
- 100 80
100
40
86 67 80
14 100
60
?7
40
20
14
57 33
14 33 20
71 67 40
67 20
57
40
67
29 67
29
86
14
86
29

100
100
67
67
67

20
20
20

43
20
14

86

22
9
9 11
9 11
36 44
18 33
45 44
9
9
27
9

44
22

11
55 22
64 56
14 36 33
43 27 89
86 55 44
71 36 67
57 55 89
57 73 100
86 91 100
86 100 78
100 67
100 27 56
71
27 33
29
82 89
82 78
14 73 67
71
36 89
71
27 22
18 33
43
27 22
73 67
43
57
18 33
9 22
14
45 67
43 45 44
100 9 11
14 73 67
33
29
14 27 33
9 11
29
45 56
14
100 27 44
43 36 33
45 67
9 11
43
14 27 11
14 45 11
14 55 78
44
14
14
14

14 9
43 18
43
29
14
14 27
29
71

44
56

11

Be

6
18
24
6
18
18
59
24

6
76
24
24
12
18
35
18
53
53
88
76
65
71
53
24
82
65
71
53
65
76
82
29
35
71
59
24
29
12
35
35
41
53
6
18
47
47
18
24
24
24
47
35
18
35
6
12

Tb

Pj

RR P-R

12
6
6
44
6

17
75
61
17
25
42
14
42
28
6
3
3
8
39
17
17
3
17
17
44
44
19
19
53
53
61
61

7
64
71
7
7
14

6
62
56
6
6
31
88
38
19

14
79
7
7

14
31
43
25
14
56
29
75
100
56
88 97 100
94 100 100
62 75 86
75
67 79
47 43
38
44
14 21
94
19 43
39 50
6?
44
19 36
6 14
38
31 43
38
94
39 50
44 21
62
31
22 29
19 14
25
31 50
62
14 7
62
19
12
50
50
31
44
44
31
25
12
25
31
6
19
12
6
19
25
19
6

6
12
24

50
36
14
7

19
12

6
3
3
19
3

14

14

14
3
6

Cp

Ce

10 53 42 50
70 87 100 100
50 80 100 81
10 ?0 25 27
8
15
27
17
50
19
31
10 7
60 47 50 38
20 60
10
10 207
o
8
19
20 20
60 73 67 31
8
20
8
20
10
30
69
50 33
8
10 47 17
70 80 58 77
80 100 83 88
100 93 100 92
100 93 100 96
80 93 100 81
60 73 75 73
46
60 20
38
10 20
19
40 53
40 87 75 38
50 27 50 23
4
8
10
27
40 13
70 53 50 27
30 20 25 19
8 38
10 47
17 23
20
23
40
8
12
19
4
4
4
4
10 13
10
10
10

6
3
3
6

Ce

10

75
TYPE
DOMINANCE

SPECIES

Amphilophturn paniculatum
vit?folium
cumanense

Cochlospermum
Erythroxylum
Calliandra
magdalenae
fendleri
Strychnos
Marsdenia
condensiflora

Belenc i ta nemorosa
Capparis indica
Platymiscium polystachium
Acacia macracantha
Lantana c?mara
Platymiscium diadelphum
Lonchocarpus atropurpureus
Simiraklu?ii
Humboldt iel la arb?rea
Helietta pleeana
spp
Ruprecht ia
Bursera simaruba
Casearia zyzyphoides
Acacia glomerosa
Macfadyena unguis-cati
Acacia pan iculata
Machaerium spp
Guapira ferruginea
Pithecellobium
ligustrinum
Marsdenia

altissima

EE

Zp

PP

-67
--3S--9--6
--33
-

47
27
40
20
47
27
13
20
27
20
60
27
87
87
73
73
33
13

57
14
57
14
86
29
43
57
29
57
71
57
71
57
57
14
14

13
7

29
29

Pithecellobium
tortum
Pithecellobium
carabobense
Guazuma
ulmifolia
Call iandra
affinis
orinocense
Erythroxylum

7
13
27
-

71
14
29
-

Machaerium arboreum
Apoplanesia cryptopetala
Capparis verrucosa
Bauhinia guianensis
Achatocarpus nigricans
Cal 1 iandra tergemina
Capparis pachaca
Coceo!oba padiformis
Paul 1 inia pinnata
Abutil?n giganteum
nov

#4

73
87
27
7
27
53
33
27

-57
-

Ch

---11-6
20
40

33
33
33
33
33
33
100
67
67
100
33
67
33
67
100
100
100
100
67

20
57.
14
20
-14 20
80
14
60
60
43
80
14
29
80
14
20
6029
29
60
14
100
100
43
14
60
80
14
40
-

--11

-----6-3
33
33
40

67-----

14
14
71
86
43
43
57
29
57
71

100
100
67
33
100
67
67
-

40

20
80
80
40
20
40
20
20
60
60-

29-

43-

14
14
14
-

14

33

Caesalpinia granadillo
Amyris ignea
Seguieria americana

20
27
40

29
43

20
60
80

47

86

67
100
67-

40

29

Casearia

praecox

20

33

60

29

Capparis
Maytenus

frondosa
spp

73
47

33
67

60
40

Eugenia

sp

Machaerium

robiniaefolium

Call iandra riparia

Bauhinia
emarginata
Randia
dioica
Casearia
aculeata
Cordia
thaisiana
culantrillo
Zanthoxylum
Randia
venezuelensis
Piptadenia
pittieri
violaceus
Lonchocarpus
Bauhinia
cumanensis

Schaefferia

frutescens

Roche-fortia

spinosa
varoasii

Aspidosperma

Euphorbia cot in ifol ia

Bunchosia

mollis

Guapira
Maytenus

pacurero
karstenii

Piptadenia

spec

iosa

Eugenia spp
sp
Chrysophyllum
Brownea
aroensis
Adelia
ricinella
Call iandra
qrac i lis
Uitex
compressa
Crot?n
niveus
Zamia muricata
Pachyptera

hymenaea

33
33

67

7-

Ximenia americana
Trichilia
trifolia

Ac

733
33
20
27
27
-20

7
7
27

14
57
86

-67
-

80

33
71
----11
29--14--6
43
33
20
33
14
40
-20
--20
40
67
40
-67

14
-

33

-33

14

13
14
13
14
20
14
----29

80

71

27
40
7
20
13
-

57
43
-

60
?7

43
57

14
14
-

33-

14
-9-6
-

14
-

---12
---18
14
14
-9-6

20

14

20
40

14
43

60
20

91112
6 6
-

60

33
--911
-20
33
20
33
60
67
20
67
60
-------------------------

11

9
9
-

11
22
-12
22
11
44
-

Be

Tb

6
18

12
6

18
18

19
12
6
12
12
19
6
31
6
6
12
19
6
6
6
12
6

18
-

22
22
11

12

36
36

22
44

24
6

6-

9
6

33

11
-

9
18
36
27
27
36
2727
-

11
11
11
22
11
33-

18
27
55
27
IB
64
9
45
27

22
22
22
11
44
11
11
33

11

18
-

11

18

11

12
6

12
6

--6

6
12

14
--11
-

------ -

36

11

-9
----------------

11

----- ----- ----- ----- ----- -----

-4

3
-

-----

--------- -

------

------

------

------------------

------------

-------

---12
-12
11

3
3
3
3
3
3
3

------

11

Ce

21

12

33
-

Cp

--6
--6

Ce

---------

38
12
-----------------------------------------

?~

11
-

6
-----12

24
29
41
12
12
6
35
6
12
12
24
6
24

RR P-R

Pj

41
29
41
18
41
12
53
29
12
29
12
18
24
29
12
12

18
9
-

---9-6
---27

100

Pd

-------

--------

-----------

76
in at least nine
however,
than
50 %.
is less

15 dominance-types;
them their presence

of

Seasonal
Beard's
Four

system

series of

are present:
rain
by decreasing

formations

series

(controlled
fall), the dry evergreen series (determined by edaph
the montane
series (controlled by
ic water deficit),
and the swamp series (controlled by in
altitude),
creasing edaphic water excess). However, not all the
in each series are represented
here.
formations
There

are other

been described

in the folded map

is shown

(see

end of this issue).

seasonal

Evergreen

soils,
ever
for

seasonal

semi-evergreen

thorn woodland,
sea
secondary deciduous
forest,

thorn

and

woodland,

shrub.

The

forest.

evergreen

seasonal

is a three-storeyed
forest with a closed tree
layer between 6 and 12 m high, overtopped
by scat
tered trees reaching 30 m. The lower storey, be
tween

variants

forest,

cactus

secondary

system. Also some

are sufficiently
established
in the series. The distribution

which

seasonal

seasonal
est, deciduous
cactus scrub, and desert;
sonal
deciduous
forest,

forest

edaphic
before in Beard's

of the formations

series

seasonal

green

had not

formations

anthropogenic
to deserve inclusion

Formation

series, located on well drained

includes 6 lowland formations and 3 variants:

The

of formation-series

of Beard's

the seasonal

of the formations

Description

2 and 5 m, is discontinuous
and includes
small trees and shrubs (Table 4). In the uppermost
tree stratum deciduous
species, with compound
crowns predominate.
leaves and umbrella-shaped
other two strata are formed mainly
by ever
are
Leaves
green mesophyllous
species.
frequently

The
between

Relationships

the three systems

of

compound
and dark

classification
association
though there is not a one-to-one
between
in the physiognomic
the categories
and
a
floristic classification
correla
(Fig. 5),
significant
tion between them has been detected by Kendall's
correlation
coefficient
z = 9.59). The
(T=0.45;
were ranked ac
classes within each classification

Even

cording to their structural and floristic complexity,

in order to perform the test (Rand, 1971). The lack
of a complete association may be due to the fact
that some of the physiognomy-types
represent serai
or
stages
types derived from
degraded community
a single association.
In some cases there is a high
of
i.e. all the evergreen thorny
association;
degree
all
scrubs belong to the C. erecta dominance-type;
the closed
juliflora

evergreen

thorny

forests belong

dominance-type.
is represented

dominance-type

physiognomy-type.
There is also a lack of a one-to-one
between

to the P.

each
However,
more
one
than
by

and

association

formation-types,

physiognomy-types
as well as between these and the dominance-types
(Fig. 5). This reinforces the evidence that the vege
tation

is represented
pattern
stages and degrees of human

by a gamut
disturbance.

of

serai

growth

in the middle

green
is formed

in the

strata and
lower

simple, thin
strata. The under

by small shrubs and coarse

forbs

Cla
{Justicia, Hellconia,
Aphelandra,
Steriphoma,
vija, Canna), but the ground is almost bare. Lianas
are present, but not too abundant.
a
is
Occasionally,
large tree, up to 3 m in diameter
found. All the stands belong to the Eugenia domi
and epiphytes

and it is one of the species-richest

for
nance-type,
mations
(Table 3).
The evergreen seasonal forest occurs from 200 to
420 m alt. on the low coastal hills, and up to 900 m
on the eastern Falcon
organic. Clearing
land has reduced

Ranges. Soils are deep and

for the establishment
of pasture
the forest area in the last decades

at present only remnant patches are left in

some areas, especially at the south of the Maritime
Valleys. Selective timber harvesting or clearance of
and

for cattle raising or for the estab

the undergrowth
have degraded
the
lishment of coffee plantations
structure to one or two layers with a tendency to
deciduousness

the tree
in the lower strata. However,
as
of the upper storey is maintained,

composition
well as some structural
column

features, such as absence

of epiphytes and palms,

cacti, presence
evidence of their being derived

plex forests.

from the more

of
as

com

D-T

P-T

Fig.
Table

5. Relationship

between

the three classification

forest
(F): Cloud
deciduous
Secondary

2. Formations

(CF); Evergreen

systems.
seasonal

Physiognomy-types
forest

77

(PH):

given

in Fig.

seasonal
(ESF); Semi-evergreen
Thorn woodland
(TW); Deciduous

3. Dominance-types
forest

(DT): as in
season

(S-ESF); Deciduous
thorn woodland
(DTW);

Cactus
seasonal
forest (Sec. DSF);
(DSF);
cactus scrub (Sec. CS); Desert
forest (GF); Dry evergreen bushland
desert (ED);
(D); Gallery
(CS); Secondary
(DEB); Edaphic
on the rock pavement
strand vegetation
(VRP); Herbaceous
(HSV). The figures indicate the number of stands in each system
Vegetation
al forest
scrub

assigned

to the next

type.

78

Semi-evergreen

seasonal

of

classified

communities

Within

forest.
as

the group

layer, between

season

umbrella-shaped

semi-evergreen

to Beard's descrip
al forest, only three correspond
or
more
less
disturbed com
tion; the rest includes
which,

munities,

formation-type.

Table

3. Number

trees

per

species

in each

family

predominate.

shrubs

Myrtaceous

are

abun

dant, but there are few herbs and there is no ground

cover. Lianas are abundant but epiphytes are scarce

however, retain elements of this

The canopy is formed by the upper

of woody

18 and 25 m, mostly
of deciduous,
trees. In the lower layer evergreen

(Table 4).

formation.

Formations

labeled

as

in Fig.

5.

Formations

Family

S-E Sec

CF
Acanthaceae

5 3 6 2

Actinidaceae

ESF

SF

DSF

Sec

DSF

TW

31

DTW

CS

CS

GF

DEB

ED

VRP

1 1 1

Achatocarpaceae
Anacardiaceae

1 112

13
1

Annonaceae

Apocynaceae

1
Aquifoliaceae
4 1 1
Araceae
2 1

Araliaceae

1 1 11

114 3

Asclepiadaceae
Asteraceae
5 2 3 3

2119
3 12 41 1

21515
Bignoniaceae
4 3 2 1

Bombacaceae

21 21 1

2565332
Boraginaceae
2 133

Burseraceae
Buxaceae

12

14

Cactaceae
Caesalpinoideae

44

1216

653644 5 6 3
158

1012

135

8
6

5 444336
1321 13

1
11
5114

Celastraceae
1

Chloranthaceae
1

Clethraceae

1 1

Cochlospermaceae
1 1

Combretaceae

Convolvulaceae

1 11
11 11111

Cyatheaceae

1 1 1

Cycadaceae

Dioscoriaceae

Elaeocarpaceae
3

Erythroxylaceae

213
1

Euphorbiaceae
Faboideae

14
1012
146

Flacourtiaceae
Gesneriaceae
Guttiferae

Hippocrataceae

6 3 2
1 11111
1 1

23
23

198
166
1 32

Hernandiaceae

41

44444

44

Capparaceae
Caricaceae

Ericaceae

21 2

56

15

79

Table

3. Continued.

Formations

Family

CF
Lauraceae

ESF

12 11

Loasaceae

Lythraceae

Malvaceae

Sec
CS

GF

DEB

ED

VRP

1
5

Meliaceae 5
Mimosoideae

6
11

2
25

6
4

9
6

10

Moraceae

111
2

Melastomataceae

1
27

1
8

13

1
4

35344

Myrsiniaceae

Myrtaceae 6
Nyctaginaceae 2
Olacaceae 2 2
7
1

Phytolaccaceae

1
1

Polemoniaceae

2 2

Polygalaceae

Polygonaceae 4 5

11

Proteaceae

Rhamnaceae 2
Rosaceae

1
2

1
3

Plumbaginaceae

5
5
1

10

Piperaceae

13 2

11

11

Rubiaceae
19
Rutaceae 4
Sapindaceae 3
12
Sapotaceae

10
8
11
1

15
11
8

10
7
8

4
5
2

12
11

Simaroubaceae

Solanaceae 4
Sterculiaceae 13 3

2
4
1
11

3
1

1111
1
1

11111

11

6
2

1
11

1111111

114

1
1

Symplocaceae
1

1232 21

Theophrastaceae
1
Tiliaceae

11212
1

11

1 1

111

12

1
2

12

1 1

Verbenaceae 3
Vitaceae
Vochysiaceae

CS

Marcgraviaceae

Urticaceae

DTW

44911322

Malpighiaceae

Ulmaceae

TW

1111

Loganiaceae

Turneraceae

Sec
DSF

Liliaceae

Theaceae

DSF

Lecythidaceae

Palmae

S-E
SF

2122122

1
21122

Zygophyllaceae

202
Species total
Genera total 136
Families total 63

12
211
149
57

292
170
55

236
135
50

2
93
62
20

87
63
34

51
40
22

1
41
32
21

51
43
26

24
20
13

49
41
24

45
35
19

38
30
19

11
11
8

80
cover degree of
from the sum of the mean
form spectra for the formations.
The relative cover degree was obtained
4. Growth
to each growth
to the total cover degree for the stand. Growth
forms: Mes = large trees (higher
form in relation
the species belonging
R = rosettes; L, s; g; H; E; P; En; h; SC and sa as in Fig. 3.
than 5 m); Mi = small trees (2 to 5 m high); N = shrubs; HI = suffructicose;
=
=
=
cover degree <5%.
+
evergreen;
deciduous;
(sv)
(d)

Table

on

<u

Mes (sv)
Mes (d)
Mi (sv)
Mi (d)
N (sv)
N (d)

35
34
+
13
33
28
+
+
10
+
+
+
00 SC

11
39
515
8
6+
15
8
+

sa

00

HL

5+

55

+
16

27+5

1111

++5

35

+0

10++++0

00

++

L
+10
En
E

7+
00

There
mation:
lum

He

Eugenia

+
7
56
5
+
+
+6+32

0
5
5
70
0

+
+
13
48
19
+

+
52
5
10
12
12

16

16

0+

7+

19

28+
+

0+
+++

+10

1 IS

+
+
12
20
13
+

00000000000
+

++

++

0+

++

000

are four dominance-types

spp, Pilocarpus

pterota-Machaerium

0
+
15
14
53
+
+

0
15
11
16
16

+++

++

000
+
19
16
7
-

?1 1

6+++0+

++

00
+
+
41
12
11
43
10
15
7
5
1113

spp-Eugenia

this for

spp, Zanthoxy
spp,

and

A.

The

Z.

pterota-Machaerium

located

0+

0++

+++0+

within

spp-Eugenia

spp

from 60 to

in the Maritime

type,
Valleys
100 m alt. and on the foothills of the eastern

Fal

The Eugenia
type is the most com
community
and
many of the spe
richest;
however,
species
plex
cies found in the Eugenia evergreen seasonal forests

con Ranges between 20 and 700 m, is not as rich in

species, but structurally complex, more mesic than
the Eugenia
forests but less xeric than the A. cryp
communities.
topetala
They may have some of the

are lacking

species

cryptopetala.

in the Eugenia

semi-evergreen

seasonal

forest.

The Pilocarpus
dary communities

woodlands

and forests are secon

from the Eugenia

forests
The
abandonment.
subsequent

derived

after clearing and

dominant
species forms almost pure stands and
tree height is remarkably uniform. Many of the un
to both. They oc
dergrowth
species are common
cupy small
the Eugenia

tracts of

land in the neighbourhood

of
forests between 400 and
semi-evergreen

800 m altitude.

found

in the Eugenia

forests

in the under

growth. Other frequent species in the undergrowth

are Bromelia
humilis, B. chrysantha, Acanthocer
eus pentagonus,
Alternan
Monvillea
smithiana,
thera williamsii.
type, the most xeric type
cryptopetala
a transition
to
constitutes
this formation,
the deciduous
seasonal forest. These com

The A.
within
wards
munities

are

located

to the west

in the Falcon

Ranges Province, between 300 and 800 m alt. The

structure is less complex,
the canopy is lower than

81

is
M. smithiana
two dominance-types.
hir
Solanum
ruscifolia,
present. Lasiacis

bare, with
for scattered colonies

ground

always

except
most
disturbed

tum,

Brachiaria
mutisii,
fasciculata,
schiediana, Elvira biflora are also found

Alseis

Acalypha
in the undergrowth.
as Oncidium

are some orchids, such

hum
and Schomburgkia

There

cebolleta

boldtii.
seasonal forest is the richest
The semi-evergreen
in woody species (Table 3). It has been greatly dis
turbed in the Maritime
Valleys and at lower alti
tudes in the Mountain
Ranges for the establishment
To the east, the forests appear as
of pastureland.
patches within a sabanna matrix and the size of the
patches has been decreasing
the last decades. The forest
fast pace by the natural
Chloris
inflata, Paspalum

continuously
during
is being replaced at a
molle,
species Panicum

virgatum, Andropogon
bicornis, or by introduced grasses such as Panicum
maximum, P. purpurascens,
Cynodon plectostichi
Cenchrus ciliaris, and
um, Pennisetum purpureum,
decumbens.

Digitada
formed
which

is per
deforestation
as the sloping terrain,
serious problems of soil ero
The

on the flat as well

is producing

sion. Frequently
the highest trees are kept in the
the wood
and
after abandonment,
pastureland
to the semi
to belong
lands are recognized
evergreen seasonal forest from their presence. The
forests located at higher altitudes have been im
poverished by timber harvesting.
Deciduous

seasonal forest. The deciduous

seasonal
in Falcon
formation
extensive
forest, the most
State, comprises 55 stands classified as non-thorny
forest or woodlands,
distributed all over
deciduous
the State. There

5 to 10m high,
In some communities

is one tree stratum

which may be closed or open.

there are occasional
emergent trees reaching up to
12m. There are no large trees. There are but few
lianas and the epiphytes
are restricted
to small
xerophytic
palms and

buttressed
bromeliads;
trees, ferns,
stilt roots are absent but column cacti

are conspicuous. Most of the tree species are decid

uous; the few evergreen species are sclerophyllous.
of the species have compound
leaves and
Many
there is a tendency to microphylly.
An understorey
of shrubs and semi-shrubs
becomes

denser

is usually found, which

as the tree coverage decreases. The

few forbs and grasses,

In the
of bromeliads.

is almost

in the other

is
the undergrowth
communities
formed by a dense layer of shrubby cacti.
The most
within
the
typical dominance-type
deciduous
seasonal forest is Bourreria cumanensis

brasiliensis-Bulnesia
Phyllostylon
arb?rea, which
occurs in the Coastal Plains, the Falcon Ranges and
the Coastal

Piedmont

constitutes

a transition

seasonal

forest

and

10 and 600 m alt. It

between
between
the

the semi-evergreen

next

type, comprising
The latter, which
billbergii communities.
lie toward the xeric end of the gradient within the
deciduous
seasonal forest, are to be found in all the

Tabebuia

physiographic
provinces.
The undergrowth
is heterogeneous,
it may

billbergii dominance-type
ther by Lippia spp, Sida aggregata,
C.

wentiana,

Opuntia

or

erecta,

and in the T

be dominated

ei

Crot?n flav ens,

Cordia

curassavi

ca. Either

this last species or Bastard?a viscosa may

be an undergrowth dominant
in the B. cumanensis
P.

brasiliensis-B.

arb?rea

forests.

In both

domi

nance-types, B. chrysantha or B. humilis may form

extensive colonies that cover the ground. Other spe
cies present in the lower layer include Cnidoscolus
urens,
M.

caesius,
gossypifolia, Melocactus
Melochia
tomentosa,
Sporobolus
Solanum
ci
argillicolum,
Eragrostis

Jatropha

smithiana,

pyramidatus,
lia ris, A. pentagonus,

Ocimum

micranthum,

Wede

lia parviflora. Most of the undergrowth

species do
not have a high presence percentage
in any domi
and they appear in patches of differing
nance-type
species composition.
To the east, the deciduous
seasonal
forest is
of
stands
Crot?n
located
represented by
heliaster,
in the Maritime Valleys and in the eastern extremity
of the Alineaci?n

in patches within
Septentrional,
the pastureland. They are all secondary forests and
in which C. heliaster behaves as an in
woodlands
vader. They are characterized
dense B. humilis or Aechmea

b*y the presence of

colonies
aquilega

loca
forming the ground layer. Their geographical
area of the Z. pterota
tion within the distributional
Machaerium

spp forests could suggest

spp-Eugenia
that the former are derived from the latter after

seasonal for
clearing of the original semi-evergreen
est and subsequent
abandonment;
however, there

82
are no evidences

to support this hypothesis.

The
within
the
and the ab
patchy pattern
pasturelands
sence of the species belonging
to the semi-evergreen
to suggest that
their former community

forests makes

seasonal

it difficult

they can evolve towards

situation
type. A different
heliaster

forests

occurs

with

the C.

in the western

located

Coastal

in which the accompanying

Plains,
species, as well
as the location suggest that they may have derived
from the T. billbergii deciduous
seasonal forest.
The

Pithecellobium

hastata
dulce-Capparis
between 20 and 600 m altitude,
stands, distributed
also belong to the deciduous
seasonal forest forma
tion. In the Falcon Ranges they are located on the
southern

slopes; and
the formation.

within

most

found

wentiana,

cur

the

nutrient-poor,

B.

they oc

Valleys
saline

viscosa

next

soils,

to the flood-prone
terrains. The trunks are thin and
whitish. The accompanying
evergreen tree species
are very conspicuous
during the dry season. The
frequent undergrowth
species are A. pentago
A. williamsi, M.
nus, A. halimifolia,
smithiana,
Setaria rariflora. Dense patches formed either by B.

most

or by B. chrysantha often cover the ground.

The physiognomy
reminds one of
and environment
humilis

the dry evergreen

mation
nate

series;
and

notably

the

with

forests

however,

of the dry evergreen

deciduous

community's

species

for

predomi

appearance

changes

seasonal forest for

Finally, within the deciduous
are found.
three C corlarla woodlands
mation,

tion between

in the Coastal

Plains

and

a transi

They represent
and the T bill
the thorn woodland

seasonal forest, from which they

bergii deciduous
have probably
derived. They resemble the thorn
in structure, except for the scattered tree
woodland
the thorny umbrella-shaped
layer that overtops
sea
trees, as a witness of their origin as deciduous
sonal forest. This
feature gives them the non
character. The presence of an al
thorny deciduous
most closed prickly pear cactus (O. wentiana)
layer
and of other invaders, such as Ipomoea carnea and
B. viscosa,
turbance.

are evidence

After
pasturelands.
abandonment,
pure
nearly
stands of Aspidosperma
spp follow, with a low uni
form stratum of this species, overtopped
by a scat
tered layer of this and other tree species. These
stands have been classified as secondary deciduous
seasonal forest. They are located to the west of the
Coastal

Plains
of B.

patches

of the high degree

of dis

and

next

Piedmont,
braslliensis-B.

deciduous

to

arb?rea

seasonal

forests. Their
as
to the
impoverished
compared
seasonal forests (Table 3), but they still

deciduous
small

Coastal

cumanensis-P.

and T. billbergii
flora has been

to it. Since the units are

species belonging
the possibility
exists that they will return to

condition. However, whether this hap

or
not
pens
depends on the pressure by cattle and
goat on the tree seedlings and their capacity to sur
vive browsing

and

Thorn woodland.
tensive

trampling.
The

formation

is a very ex

thorn woodland

located

in the Coastal

Plains,
it can reach the seashore, in the Coastal Pied
mont
and in the Central Depressions,
on
mostly
flat terrain. It comprises mainly
closed to desertic
woodlands
(Fig. 5); however, the vertical structure
where

is very variable
as

the seasons.

They occur to the west,

in the Coastal Piedmont.

have

their former

O.

In the Maritime

forest has been greatly

activity. Large tracts of forest
for
the
establishment
of

by human
been
felled

are

frequently

seasonal

disturbed

have

tomentosa.
lowest,

deciduous

represent a xeric extremity

The undergrowth
species

and M.
on

The

forests

to 5 m

and

high,

and some units have been classified

'cardonales'.

The

is dominated

microphyllous
near
branch

by

tree

layer,

from

evergreen
Trees
legumes.

thorny

species,
mostly
the ground
and are
(1 to 2 m)
are
There
trees
umbrella-shaped.
sclerophyllous
and shrubs, but orthophylly
both in
predominates,
and in plant cover. The under
species number
is dominated

by shrubby cacti. Ground

is very sparse and grasses are almost ab
column cacti may overtop the tree layer.
The most extensive thorn woodlands
belong to

growth

vegetation
sent. The

the P. juliflora dominance-type.

Their large exten
sion may be related to the ability of Prosopis
to ob
tain water from underground
This
storage.
species
does not show xeromorphic
features. It evades wa
ter stress by means
by the high

of a deep

rooting system, as
rates maintained
transpiration
even
the
and
the
along
day
during
dry season. The
Coastal
Plain
soils, where this species competes
shown

83

advantage, must have a large water retention

capacity and most probably upward flow from the
water table maintains
the storage in the deep soil

with

layers.

thorn woodland

The

has

of

the domain

been

and the
the Europeans
goats
is impoverished,
specially the palatable
vegetation
this vegetation
type has been con
species. Though
since

the arrival

of

'typical' of the semiarid

even a climax (Tamayo, 1967),
sidered

represents
could become

coastal

zone,

it is likely

and

that

it

which
community-type,
due to the tenacity of

secondary
established

both P. juliflora and O. wentiana, whose dissemi

nation is favored by goats. The former is dispersed
tract, where seeds are
internally via the digestive
the cactus

scarified;
the joints

carried

on

regenerates vegetatively
the goats' hide. Other

from
evi

Ritterocereous

spp

accom

The

dominance-type.

panying tree species and the undergrowth

composi
tion do not differ from those of the P. juliflora
woodlands.
The P. juliflora-Ritterocereus
spp
thorn woodland,
which represents a transition be
tween this formation and the cactus scrub, includes
woodlands
and 'cardonales'.
Both physiognomy
types differ from each other in the relative height
of
the trees and
In the
the column
cacti.
the

'cardonales',

cacti

become

very

be

conspicuous

cause

the tree layer, which

forms a
they overtop
crowns.
very uniform canopy of umbrella-shaped
In the woodlands,
the trees are as high or higher
than the cacti. In both physiognomy-types
the
cacti have a bole attaining more than a me
ter high and more than 30 cm diameter, with many
branches and are heterogeneous
with regards to
size
and
The
form,
probably age.
undergrowth may
column

to support
that the P.
dences
the suggestion
is a secondary community-type
juliflora woodland
sea
are the presence of elements of the deciduous
sonal forest in some of the stands, as well as the oc

Piedmont.

currence of patches of that formation

intermingled
with it. It appears that this group of thorn wood

In the Paraguan? Pen?nsula, three C. erecta thorn

are found. They have been classified as
woodlands

lands has been derived

from the deciduous

forest,

however,
as to be considered

a formation-type
to Opuntia,
the most
cies to be found in the undergrowth,
In addition

M.

datus,

seasonal

with Tamayo's (1963) proposal;

so
it has become
established
sufficiently

in agreement

ciliaris,
Euphorbia

caesius,

A.

urens,

Ayapana

its own.
spe

are S. pyrami

Allionia

E.

incarnata,

Caraxerum

halimifolia,
dioica,

on

common

squarrosa,

vermicularis,
B.

viscosa,

J. gossypifolia,
C. erecta and C. flavens
however,
in
the
last
four
only
species, may become dominant
the

There

thorn wood
is a variant of the P. juliflora
the dominant
species adopts a
from the
form and branches
growth

in which

shrubby
of goats brows
ground, probably as a consequence
case
the vegetation
ing the seedling apices. In this
becomes almost impenetrable,
cluttered by the tree
branches
Even
juliflora
are also

and

and differ from

evergreen thorny woodlands
the shrubs are
erecta shrublands
because
a
tree
and
they are located on
by
layer
overtopped

open

the C

cacti.

shrubby
though most of the stands belong to the P.
other dominance-types
dominance-type,
Whenever

become

represented.
abundant

juliflora,

the stand was classified

and

the column

co-dominate

cacti

with

in the P. juliflora

P.

soils.

non-calcareous

Two

of

these

oc

communities

at 120 to 200 m alt.

cur in a less xeric environment

Santa Ana hill, next to a T billbergii forest.

of
site has been cleared for the establishement

around
This

sesame,

and

crops

mainly

days

is one of the rural areas with

sorghum

density.
possibility
forest occurred at this site cannot
B.

dulce-C

cumanensis-P.

hastata

corn

and

nowa

higher human
seasonal
that a deciduous

The

The

understorey.

lands,

It oc
by a closed layer of O wentiana.
curs in the Central Depressions
and in the Coastal

be formed

be discounted.

brasiliensis-B.

and C. corlarla

stands

arb?rea,

included

P.

in

might have been

derived from the deciduous
seasonal forest since,
to the
even though their physiognomy
corresponds
former, species typical of the latter grow here.
the thorn woodland

formation

However,
they are evergreen and thorny. They are
located in the proximity of villages or pasturelands.
the thorn
Within
the seasonal formation-series,
woodland
reduction
ble 3).

represents
in woody

a stage
species

in which
number

a drastic
occurs

(Ta

84
Deciduous

thorn woodland.
not

woodland,
system,

thorn

deciduous

in Beard's

included

resembles

The

classification

in structure;
the tree
dominate

the thorn woodland

except that deciduous

species
stratum. It can be barely distinguished

growth does not differ from that of the P. juliflora

in composition
woodlands
but it is sparser. They
on

grow

non-calcareous

next

the

Cactus

scrub. The

during the wet season, but in the

season
It
its
appearance
dry
changes considerably.
is not clear whether
these communities
have der

series

is the cactus

from

thorn woodland

ived from deciduous

seasonal forests; although this

cannot
be
discarded, features to support
possibility
this view are not present, for most of the deciduous
trees

are

low

thorny umbrella-shaped
from the deciduous
seasonal

They differ
the predominance

of

legumes.
forest in

their

species composition

woodlands

forests,

and

are distributed

by

comprises

in all the Physiographic

Provinces
lands in the
region, on disturbed

the dry

neighbourhood

of villages. The dominance

of C.
be due to the fact that it is used nei

praecox may
ther as firewood nor for timber. Some time ago it
was gathered
for manufacturing
soap, but this
has
been
abandoned.
The
practise
pods are not
palatable for goats and it is one of the few species
that is not eaten by them. For these reasons C prae
cox has an advantage over other woody
species in
such semi-artificial
terocereus
Pereskia

guamacho,

and Capparis
erecta,

O.

landscapes.
A.

tortuosa,

odoratissima

wentiana

or

P. juliflora

with

codominate

may

I.

C.

Mimosa

may

the undergrowth,
whose composition
that of the P. juliflora woodlands.
The C
deciduous

and Rit
and

praecox,

arenosa

are very abundant.

carnea

erecta stands classified

thorny woodlands,
the deciduous
thorn woodland.

by cacti, forbs, ephemerals

is sparse and patchy and the ground is
bare. Even though the canopy may be decid

almost
uous,

evergreen

dominate

is similar

Of

the 21 'cardonales'

cactus

The occurrence
with

scrub

of patches of
formation

other

intermingled
types, as well as evidences of tree felling in some of
com
them, suggest that they might be secondary

or impoverished
thorn woodlands.
They
could also be remnants of larger cactus scrubs that
have been either encroached
by desert or invaded

munities

of
by P. juliflora. Most
terocereus dominance-type,

to the Rit
them belong
in which R. griseus or
The dominant
species in

R. deficiens are dominant.

the undergrowth may be O. wentiana, A. halimifo
lia, or C erecta, and the accompanying
species are
the same as those found in the P. juliflora wood
lands.

of dense, almost
'cardonales',
consisting
to
not
of
do
column
stands
cacti,
pure
correspond
because
the cactus scrub in Beard's classification,
Other

the vegetation
classified

been

of

is too dense

and high. They have

cactus
scrub. The
secondary
the
column
cacti,
together with
as

in

abundance

to

in height of the individuals, suggest that

uniformity
these communities
represent
early stages of a
succession,
taking place after abandon
secondary

as open or desertic
are also included in
Even

shrubs

among

predominate

species

trees.

is im

several dominance-types
(Fig. 5). All the Cercidium
are
in this formation. They
stands
included
praecox
within

sparse. The
formed

undergrowth,
and grasses,

in Falcon State,
present
stands
with
Beard's
of
agree
only eight
description
are
cactus
at
scrub
located
the
the
(Fig. 5). They
north of the Coastal Plains, bordering the desert or

represented

'cardonales',

shrubs, which are

total plant cover is low. The

the trees and

the thorn woodland.

poverished
(Table 3).
The deciduous
thorn woodland,

as a thorny
cacti occur

scrub, described
in which
the column

and

thorny species,
and the abundance
of shrubby cacti in the
feature is the presence of
Another
undergrowth.
column cacti, which can be very abundant
in some
(Table 4). Woody

low and

of the seasonal

formation

lower

canopy,

stands

vegetation-type
scattered among

soils.

the

though
shrub dominates,
the degree of cover of the accom
panying trees is higher than that of C erecta and
the structure and appearance
differs considerably
from that of the C. erecta shrublands. The under

ment

of

cleared

tracts

of

and
germination
structure
terocereus.
The

chronized

it cannot

is not

be considered

cluttered

and

between

the thorn woodland

the mesic

syn
land, through
establishment
of Rit

to xeric climatic

but
simpler,
intermediate

and the desert

gradient.

The

along
cacti are

85
small and branch very little and near the ground,
attest to their recent origin. They all belong

which

to the Ritterocereus
and

deciduous

mostly

ties are reduced

of the Coastal

and are

spp dominance-type,
The

non-thorny.

in extent and

Plains,

located

in the Coastal

communi

to the south

Piedmont

and

the decidu

in the Central

bordering
Depressions,
ous seasonal forest, from which they have probably
se
derived. They belong to the seasonal formation
ries since they grow on well drained soils. Their
high species richness as compared
scrub (Table 3), is another evidence

to the cactus
of their higher

complexity.

Dry evergreen bushland. The dry evergreen bush

land consists of a low layer (0.5 to 2 m high), domi
trees and bushes,
nated by evergreen sclerophyllous
with a tendency to microphylly,
overtopped by scat
tered trees and column cacti. Thorny
species pre
dominate. Ground vegetation
is remarkably scarce,
shrubs show a
sclerophyllous
no
with
obvious
trunk,
habit,
hemispherical
growth
and the landscape appears greyish-green due to the
grasses

(Table 4). The

whitish

branches

10 of

Only

the 46 desertic

physiognomy
within
formation-type

types represent the desert

the seasonal formation
series (Fig. 5). They are lo
cated on the northern fringe of the Coastal Plains.

and the light green foliage. Plant

less than 50% and the soil
of the stands belong to the

cover

is low, sometimes
surface is exposed. Most
erecta

C.
Desert.

and a few forbs and

of small Cactaceae

consisting

In the undergrowth,

dominance-type.

Malvastrum

A.

americanum,

A.

incarnata,

squar

and A. pittieri may be

adscensionis
rosa, Aristida
come abundant, but the most frequent species are
M.

and

caesius

O.

wentiana.

The desert appears either as scattered herbs, shrubs

and low trees on a bare plain or as patches of vege
tation complexes scattered on a bare plain. In both

The dry evergreen bushland occurs in the Coastal

Plains, at 20 to 90 m alt. and in the Central Depres

cases P. juliflora
is the most abundant species. The
are
small (2 to 3 m in diameter) and rather
patches
dense. The vegetation within the patches resembles
a thorn woodland;
however trees are dwarfed and

however it
to the thorn woodland;
close proximity
structure
in
and
in
differs both
species composi
are caused by the soils, which
tion. The differences

with

to the leeward due

their branches

to

swept
action, and the flora is poorer (Table 3). The
density of patches, as well as their size and the den

wind

to
sity of vegetation within the patches decreases
ward the shore untill a point is reached where they
to the
contain dead trees. This fact, in addition
presence

of

remnants

within

of

deciduous

seasonal

forests

suggests that this may be

It is unlikely that vegeta
itself, for much of the exposed

the desert matrix,

a secondary
formation.
tion will re-establish
soil has probably

been removed by wind

The dry evergreen formation

and water.

series

are

structure
by

the

and physiognomy
limiting moisture

quently
terrain

or

soil

posed by topographic
override the effect of rainfall

conditions,

which
Two for

seasonality.
are represented
mations
in Falcon:
the dry ever
on the rock
and the vegetation
green bushland
pavement.

and

by gravel or pebbles.
the

emphasizes

lies in

It often

alt.

calcareous

overdrained,

overlain
surface

fre

The whitish
appear

landscape's

ance. Forty-six species in 36 genera and 23 families

have been listed (Table 3).
The

Rock pavement.
ment

of

consists

vegetation
scattered

low,

on the rock pave

shrubs,

herbs

and

in the central
cacti growing on a desert pavement,
in
the Central
Plains
and
the
Coastal
of
portion
are
tree
entire
The
almost
few
species
Depressions.
to the dried water courses. The land
ly confined
scape is dominatd
by the greyish sclerophyllous
cacti outstanding
and the low globose
bushes
against

are largely
supply im

and

sandy

cies

Vegetation
determined

180 and 200 m

sion between

the desert varnish. The most

are C.

ment

erecta

and M.

and C praecox,

and Haematoxylon

caesius,

on

abundant
the

rock

spe
pave

P. juliflora, C odoratissima
the water
brasiletto
along

courses.

The narrow

fringe of littoral hedge, located be

and the
tween the strand herbaceous
vegetation
in the isthmus
thorn woodland
along the coastline
and

the eastern

shore

of

the peninsula,

should

86
probably be included in the dry evergreen forma
tion series. This area is subjected to the winds load
ed with salt spray that come from the sea. The
dominant

erectus,
grows
it fixes with its

Conocarpus
species,
on the sand dunes which

stunted

system
extended

rooting
branches

cies present

on

in the wind direction.

its

and

Other

spe

are Sporobolus

virginicus, Egletespros
The tracts of
argothamnoides.

trata, Argythamnia
littoral hedge are smaller
and have

side

the windward

not

field

been

area

than the minimum

in the present

surveyed

study.

Montane

All

stays soft and humid all year round.

to the Eugenia
the stands belong
become

series cannot

in Fal
be represented
on the one hand, the lowlands

complete
con State because,
are dry and thus the gradient goes upwards from
xeric to humid vegetation
types. On the other hand,
are not high enough to attain the
the mountains
conditions
ecological
of the series develop.

in which
There

dominated
paraguanen
by Geonoma
community
a
area
at
to
summit of
the
small
is
confined
sls,
hill (850 m) on the Peninsula. Here, the
adopts a dwarf habit, probably
due to the prevailing
strong winds.
Below the palm brake, between 700 and 800 m
Santa Ana

alt.,

there

species

is a mountain

rain

forest;

the

however,

be

is located
largest extension of this formation
tween 1000 and 1 500 m alt. on the eastern extremi
range. There are small
ty of the northerly mountain
patches of cloud forest scattered on the summits of
the other ranges too. These represent remnants left
on the least accessible

Piperaceae

prominent

the former

families,

tresses, but

trunk diameter

evergreen

seasonal

dominate.

The

forest.

is smaller
Evergreen

than

in the

species

pre

species list includes 202 identified

in 136 genera and 63 families; how
be underestimated
since it has not

woody species,
ever, these may
to identify all the specimens collect
been possible
ed. Some of the species found only in this forma
petiolaris,
M.

aeruginosa,

P. perquadrangularis,

racemosa,

Gonzalagunia

costanen
dicocca, Rapanea ferruginea,
Psychotrla
Plukenettia
sp, Pleurothallis
sis, P. guadalupensls,
camensis,

Evodlantus

E.

heterodoxum,

Swamp

Formation

tum,

funifer,
E.

Epldendrum

lividum,

and

many

elonga
more.

the last formations

are only two montane

area:
the palm brake and
in the study
formations
rain forest. The former, a reduced
the mountain

dominant

spp
and

the higher
among
conspicuous
being
specially
trees. Dark green thick simple mesophyllous
leaves
are
trees
There
with
few
plank but
predominate.

Miconia

The

Lauraceae,

dominance-type.

Moraceae

tion are Palicourea

series

Formation

and

sites.

forests have three layers, with two tree layers

at 14 to 20 m and at 10 to 12 m high. The third lay
er, formed by shrubs, palms, ferns and small trees,
feature is the
reaches up to 2 m. The outstanding
The

series

wood
is represented by the mangrove
in the
lands, of which two types can be recognized
occur
area:
the
the
that
forests,
along
bays
study
and the coast within the Maritime Valleys Province,
and the shrubby patches of the arid zone along the

This

series

western

and
the Paraguan?
Pen?nsula
forests
de Coro. The mangrove
are three storeyed, with the closed canopy at 20 to
26 m high, an open layer at 10 to 18m and another
closed stratum at 3 m. There are few shrubs, epi
phytes or lianas. Palms may be present. Towards the
around

coast

of

the Golfete

their height and floristic and structural

tree species
decreases.
The dominant
complexity
are Rhizophora
Avicennia
and
mangle,
germinans
seashore

Laguncularia

racemosa.

great

are restricted
The arid zone mangrove woodlands
to a few spots along the coast. They are extremely
simple in structure. They consist of almost pure
1 to 3 m high, growing
stands of A. germinans

whole

small ponds and tidal inlets and surround

ed by halophytic
herbaceous
vegetation, mainly of
ritteriana
Sesuvium portulacastrum,
Heterostachys

which
of
among
epiphytes,
profusion
mosses
liverworts
and
ferns,
orchids,
bromeliads,
are found on tree trunks and branches through the
rounded

profile.

Lianas

and small. The

are

rare,

tree

crowns

soil is dark brown,

are

organic

around

and Salicornia

sp. It is hard

to say whether

these

87

are

communities

remnants

of

larger

or

one

gallery forests occur in the Coastal Plains

Central
the rivers and
Depression,
along

The

early

It has
woodlands.
stages of developing mangrove
been suggested that the whole Golfete de Coro was
forest; however, there are
occupied by a mangrove

and

no evidences

long to the P. juliflora

dominant
species adopts a tree habit, with a 5 to
8 m bole and a very extended
umbrella-shaped

The presence
to test this hypothesis.
of a forest was not reported by Americo
Vespucci
in his description of the vegetation of the Venezue
lan coast, along which he travelled with Ojeda
Surinam to the Venezuelan Gulf in 1499.

from

to the mangrove
woodlands,
submerged
are
of the vascular plant Thalasia
communities
are very rich habitats, where many
marine species in their larval stages strive. Among
and Sar
the algae, Gracilaria, Gelidiela, Halimeda
These

are

gassum

Other

not considered

in

strand vegetation,
the gallery forest and the edaphic desert, which lie
and are
side by side with the seasonal formations
conditions.

by edaphic
The herbaceous
strand communities
eastern

coast
have

they

of

the Paraguan?
classified

been

occur along
Pen?nsula,
as savanna

(Tamayo, 1941) and in small patches along the in

land coastal fringe. The formation comprises a het
in which floristic
erogeneous group of communities
gradients perpendicular
seaward extremity
The
phytes, such as Mallatonia
H.

tulacastrum,

to the shoreline

are found.

is dominated

by halo
S. por

gnaphaloides,
Salicornia

ritteriana,

spp,

among

reaches true dominance.

others, none of which
Semishrubs and small shrubs with a prostrate hab
it,

such

as E.

cum, Crot?n
cistoides,
grasses
Cenchrus

prostrata,

are

thorny

at 8 to 10m. They be
Here, the
dominance-type.

very

conspicuous

and

Arrabidea

with P. juliflora. O
frequent species in the under
it is not abundant,
it is con

codominate

may
is the most

wentiana

and, though
spicuous in the otherwise sparse species poor lower
Ruellia and Cardiospermum
layer. S. argillicolum,

growth

areas occupied
cactus scrub,
by thorn woodland,
and desert, where its width has been exaggerated

There are three community-types,

Beard's system: the herbaceous

the

Lianas

evergreen

storeyed

canopy

in the understorey.
frequent
Forty-nine
in
listed, distributed
woody
species have been
41 genera and 24 families (Table 3).
The gallery forest is particularly noticeable
in the

edaphic formations

where

one

a closed

are also

abundant.

determined

crown.

are

They

mollissima

Next

found.

streams.

forests with

Heliotropium

curassavi

Tribulus

however,
pilosus);
these communities

they grow very sparse

cannot be considered

vermicularis,
ovalifolius,
occupy the continental
extremity. Some
are present (S. pyramldatus,
S. virginicus,

ly; thus,
The sandy soils
savannas, since forbs predominate.
and the winds are the limiting factors. In recent

is
1960, 1966). Even though this formation
law
it
del Ambiente),
protected by
(Ley Org?nica
has been greatly disturbed for the establishment
of
(Hueck,

and

cropland

it has disappeared

(Pia, 1980).
The edaphic desert
4 m)
of
scattered

at localized

spots

of a low layer (2 to
deciduous
mainly

consists
trees,

mierophyllous
legumes; evergreen shrubs predomi
nate in the lower stratum. Semishrubs,
forbs and
grasses grow sparsely. The ground is almost bare,
except

for the small cactus M.

though
ble
O

very

4). Other

sparse,

present

species
A.

wentiana,

becomes

incarnata,

differs from
formation-type
woodland
because vegetation
it cannot

be included

caesius which,

even

conspicuous

(Ta

are Opuntia
S.

caribaea,

pyramidatus.

This

the deciduous

thorn

is sparser and lower;

se
in the seasonal formation

ries because

aridity is determined
by the steep dis
and
the
rather than
surfaces
slopes
gravelly
by climatic conditions. This formation might have
derived from the deciduous
seasonal forest, but the
sected

to test this hypothesis.

evidence
is not sufficient
The upper tree layer is absent and the vegetation
is
scattered. The edaphic desert is sufficiently
extend
ed, stabilized and recurrent in similar environments
so as to deserve being introduced as a new category.
It occurs in the Coastal Plains,
the Coastal Pied

located in the eastern coast

years, the communities
of the Peninsula, have been gradually invaded by O
wentiana due to an increase in the goat and donkey

mont,

the Central

populations.

Ranges,

between

and the Falcon

Depressions
80 and 400 m alt. It belongs to C

88
erecta or C. corlarla dominance-types.
It is richer in
species than the seasonal desert (Table 3).

and conclusions

Discussion

That

and climate

are closely related

a
the
recognized for
long time. However,
such as those proposed
simplistic approaches
by
Holdridge
(1959) and Sarmiento
(1968), should be
taken with care. It is unfortunate
that Holdridge's
vegetation

has been

model
but
is artificial,
and to handle by a wide range
trained per
non-scientifically
It has practical
for
planners
significance

The

physiognomic
easy to understand
of users, including
sonnel.

and policy makers,

classification

as well

as for agricultural,
sil
research. It also
and ecological

vicultural, faunistic,
in a simple fast way.
allows monitoring
vegetation
em
and structural
life-form
The
categories
ployed, as well as the scales and the techniques for
their assessment

were chosen

from those

suggested

in the literature

1967;
1957; Fosberg,
(Dansereau,
& Ellenberg,
Mueller-Dombois
1974); however,
and on the
criteria based on local characteristics
were
a
to
system,
necessity
comprehensible
produce
for the data

applied

The

description.

analysis

categories,

and

the structural

terms, and techniques

and a check
defined,

employed were previously

sheet was devised and used during field work, in or
as far as possible the subjectivity,
der to minimise
studies. Special
in vegetation
which is unavoidable
care was

circular reasoning and the

and vegetation were gathered
to the in
previous
independently

taken to avoid

data on environment
and

analysed

(Matteucci & Colma,

terpretation
1982).
studies frequently
Local vegetation
consist

of
clas

1973).
1971; UNESCO,
1967; K?chler & Montoya,
This approach may lead to more subjective deci
information
valuable
sions than necessary. Also,
to unwilling oversight of details
of classes. We
for the definition

could be lost due

not

considered

round and first obtained

other way
in a way as objective as possible, producing
from the analysis of the data.
the classification
was
with
the information
gathered
However,
the

records

as to allow the in
and consistency
terms
the existing clas
in
of
of
results
terpretation
sification systems and enable comparisons with the

enough

detail

vegetation

of other

regions.

types in Falcon State should be considered

caution due to the lack of adequate climatic

with

data
than half of
(Matteucci & Colma,
1986). More
the meteorological
stations are located in towns or
or
in
lowlands and valleys and
villages,
protected
their distribution

is uneven.

?vapotranspiration

data.

Relief heterogeneity
the extrapolations
to nearby areas. The
precludes
observation
intervals are short in most cases and
a
few
of
them collect both temperature and
only
The 39 stands of natural
stations

meteorological
Sarmiento's
The

(1968) orthogonal
system
(Fig. 6a).
scrub and the desert fall within
the
scrub niche, and not in the desert and semi

thorn
desert
This

vegetation within which

are located, are plotted
in

cactus

as would have been expected.

due to the fact that rainfall seaso

environments
is probably

have a greater
nally and rate of ?vapotranspiration
influence on physiognomy
than mean annual tem
The possibility
that the
perature and precipitation.

both

scrub stands are degraded thorn

that
rejected on the grounds

and cactus

desert

woodlands

relating the stands to some previously adopted

to simplify
sification.
Keys have been prepared
to the correct category
their assignment
(Fosberg,

went

has gained favor in Venezuela,

through its
application
by Ewel & Madriz
(1968). Any associa
tion between
climatic parameters
and vegetation

may
fall within

be

Bailey's

arid Moisture

Province

(Fig. 6b).
In previous studies, the constraints
imposed by
edaphic factors have not been considered. The dry
as thorn
has been classified
evergreen bushland
or
woodland
cactus scrub (Ewel & Madriz,
1968;
Hueck,

1960, 1966; Pittier, 1937; Sarmiento,

1972,
et
Smith
re
without
1976;
al., 1973; Tamayo,
1958)
to
and environ
the distinctive
gard
physiognomic

mental

features.

In none

of the papers mentioned

It has probably

the edaphic desert is recognized.

in the deciduous
been included
though
edaphic

even

is lower and sparser. In the

is controlled
by terrain
vegetation
a distinction
that has not been mentioned
the former
desert

factors,
in reference
bility

bushland,

to the deciduous

remains

that

The possi
desert is derived

bushland.

the edaphic

89

29H

?2 o7

130

06

ft(15 #ol38e%7
8
6

2o

?0

130

%
4o
4o

70

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iR

40

4o!

?26f
UJ

4p

%25h
2'
LU

3o
1
1

3o

i24h

THORN SCRUB

';23h
<

2 221
<
UJ 85
?3
2 21 O

TROPICALFOREST

a SUBMON-, SUBTROPICALft TROPICALSUB

SUBTROPICAL
TANEDECIDUOUSSEASONj MONTANE FORESTS
AL FORESTS
lo

o
I
UJ
(O

1,
I0RNFOREST

t-*

fe iU te fe iV

is 14
ra-~n12
-8-Hr
ANNUAL PRECIPITATION (dm)
T
T

MEAN

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12

^12

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z 10
O

#7

4 4

?O 8
h e
er
UJ
CD

?I?*t3

,
2

.2
I
.2

ARID

SEMIARID

3
between
Provinces

vegetation

MOIST
SUBHUMID

9
7
8
4
5
6
BAILEY'SANNUALMOISTUREINDEX
and climate.

(6b). Subdivisions

Thirty-nine

within

stands

the graphs

HUMID
_J_L

_I_L.

-L.

_L_

Fig. 6. Relationship
to Bailey's Moisture

IDRY
?SUBHUMID

10

are plotted
in Sarmiento's
orthogonal
to the formation-type
niches

correspond

12

II

(6a) and
(6a) or moisture

model

in relation
provinces

(6b). 1)CF; 2) ESF; 3) S-ESF; 4) DSF; 6) TW; 7) DTW; 8) CS; 10)D; 12)DEB; 13)ED; 15)HSV.

from the deciduous

are no

structural

seasonal
or floristic

forest; however, there

to support
elements

the hypothesis,
neither the ability to regenerate a
ever
if
it
forest,
existed, for not only the genetic
has
but also the present soil condi
been
lost
pool
tions would
Temporal
in previous

preclude

its establishment.

have also been neglected

relationships
studies. Either vegetation
types which

in origin have been classed together or vari

ous disturbed
types have been classi
community
fied in separate groups even though their origin is
differ

the

same.

For

the deciduous
bushland
example,
et
and
al.
Smith
described by
(1973),
regarded as a
new vegetation
type by Sarmiento
(1976), is derived
from

the deciduous

has been classified

seasonal

forests

in the present

and as such

paper.

It is lower

90
than

the deciduous

forest described
seasonal
by
thorn wood
deciduous
but
than
the
Beard,
higher
land. The tree species in the upper layer belong to
the deciduous
seasonal forest, and they lie side by
the

side with
structural
however,
disturbance
son

latter.

features

It seems

are due

that

obvious

their

to assume

that

the

the soil, there is no rea

forest will not recover.

this happens will depend on the regenera

tion pathway followed by the secondary
successin
(cf. Van der Maarel,
1984).

scrub as defined by Sarmiento

(1976)
a tree layer of up to 8 m high, and does
not correspond
to Beard's description
of the cactus
cactus

includes

scrub. In the present study, the existence of such a

tree layer, as well as the geographical
location, are
considered as an indication that these cactus scrub
communities

owe their existence

to anthropogenic
on the deciduous
seasonal forest

forces operating
and have been classified as secondary
or as deciduous
seasonal forest.

cactus

scrub

is posed by Beard's (1944) and

of the thorn wood
Sarmiento's
(1976) descriptions
land. According
to the latter, the thorn woodland
has two tree layers and the upper layer attains 8 m
of height. On a close inspection it turns out that the
species in the upper tree layer are those of the
seasonal
deciduous
forest; thus, in the present
to Sarmiento's
the stands
study
conforming
have been classified as deciduous
description
sonal forest or secondary deciduous
seasonal

sea
for

est.

(1953) have recognized a new

Asprey & Robbins
formation:
the cactus-thorn
the
scrub, in which
column cacti are very conspicuous
and overtop the
tree layer. This structure is present in Falcon
State too, but it has been classified as thorn wood
closed

or secondary cac
thorn woodland
land, deciduous
on the species composition
tus scrub, depending
and the history of the site.
A

to the study of
holistic
dynamic,
approach
discloses
the
and
vegetation
temporal
spatial rela
a
tionships. However, only
thorough knowledge ob
tained through
interpretation.

intense field work permits

(eds),

J. S.,

1949. The

J. S.,
A.

this kind

of the Windward

vegetation
For. Mem.

and

192 pp.

of tropical American

vege

36: 89-100.
S. D.,

Nacional

1982. An?lisis

del

Estado

Falc?n:
Francisco

Experimental

1957. Biogeography:
York.

P.,
New

J. & Madriz,

Memoria
ciones

Ministerio

sobre

Nacional

de Agricultura

F. R.,

de la
regional
Los suelos.
de Miranda,

1967. A

Zonas

de

vida

perspective.
de Venezuela,

2da ed. Edi

ecol?gico.
de Investigaciones
Agropecuarias,
y Cr?a, Caracas.

classification

IBP Handbook

L. R.,

Holdridge,

an ecological

el mapa

In: G. F. Peterken,

IBP areas.

tions

1968.

A.,

explicativa
del Fondo

Fosberg,

Guide

of vegetation
for general
to the check sheet for

4, pp 73-120.

1959. Determination

Oxford.

Blackwell,

of world

forma

plant

from

data. Science
105: 367-368.
simple climatic
R. A.,
1973. The vegetation
of the Antilles.
In: A. Gra
and vegetational
in northern
(ed.), Vegetation
history

Howard,
ham

Latin America,
Hueck,

vestigaci?n
Hueck,

pp

1-5.

K.,

y Capacitaci?n,
1966. Die W?lder

A. W. & Montoya
of vegetation:
sification
'
21: 98-109.
,

K?chler,

J. F.,

South

1973. On

America

and

and human

environments,

zona

Tucson.

Press,

1961. Atlas

M.A.C.,

M,

Fischer,
Stuttgart.
1971. The UNESCO
clas

J. M.,

some

tests

in the tropics.

(eds), Coastal
pp 75-90.

forestal

de

y Cr?a, Direcci?n

Agricultura

Venezuela.

M?rida,

S?damerikas.

Turrialba

the origin of the dry climate

in northern
the southern
In: D. H. K.
Caribbean.

& A. W. Wilson

Amiran

Amsterdam.

Elsevier,

1960. Mapa
de la vegetaci?n
de la Rep?blica
de
Bol. 7, Instituto Forestal Latinoamericano
de In

K.,

Venezuela.

deserts:

their natural

The University

Venezuela.

de Recursos

of Ari

Ministerio
Naturales

de

Renova

Caracas.
S. D. & Colma,

Matteucci,
dio de

la vegetaci?n.

A.,

1982. Metodolog?a

S. D. & Colma,

para

el estu

No

22, Serie Biolog?a. Or

DC.
168 pp.
States, Washington,
1986. Caracterizaci?n
clim?tica
A.,

Monograf?a

of American

ganization
Matteucci,

Ecol

Falc?n.

Ronald,

bles,

Lawton

Ecological

Berlin.
Springer,
in tropical America.

y el ambiente

Dansereau,

Lahey,

environments.

classification

& Matteucci,

Universidad

Ewel,

natural

Ecology

vegetaci?n
Coro.

and dis

& H. W.

vegetation

Oxford

1955. The

types.

Colma,

their definition

Cannell

semi-arid

34, pp 73-97,

Islands.

tation

of Jamai

vegetation

127-158.

Leeward
Beard,

in

1944. Climax

25:

Beard,

climates:
G. H.

Vol.

J. S.,

ogy

1979. Semi-arid

Agriculture

Studies
Beard,

1953. The

In: A. E. Hall,

P.,

purposes.

A similar situation

of

H.

Bailey,

to human

Whether

The

R. G.,
Asprey, G. F. & Robbins,
ca. Ecol. Mon.
23: 359-411.

tribution.

interference;
since seed sources have been left and the
has not altered

References

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Falc?n. Acta Cient. Venezol.

37(1): 63-71.
S. D., Colma, A. & Pia, L., 1979a. An?lisis
regional
de la vegetaci?n
del Estado
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Falc?n: La vegeta

Matteucci,
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Publicaciones

tituto Universitario

de Departamento
de Investigaci?n,
de Tecnolog?a
de Coro, Falc?n.

Ins

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S. D., Colma, A. & Pia, L., 1979b. An?lisis

regional
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la vegetaci?n
y el ambiente
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de Departamento
Publicaciones
Geolog?a.
de Coro, Falc?n.
de Technolog?a
Instituto Universitario

Matteucci,
de

S. D.,

Matteucci,

resources

and

land mapping

1982. Desertification

L.,

Univ.

Stoffers,
Tamayo,

231-242.
D. & Ellenberg,

Mueller-Dombois,

and methods

1974. Aims

H.,

of vegetation
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naturales
Falc?n. Recursos
1958. Estado
O.T.E.E.,
de

su

econom?a

bles,
Pittier,

1920.

Venezuela,

los

en
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L.,

VII/2,

Proc.

Rand, W.,

Am?rica
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Bol.

Int. Soc.

from aerial

Photogrammetry,

J. Amer.

Cie?e.
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Nat.

F.,

seasonal

South

America.

plant

de

Bol.

Soc.

convergences

be

simples.

J. Ecol.

60:

floristic

and

formations

of

367-410.

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Rev. For. Venezol.

xer?filos

de mapa
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Bol.

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F.,

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costanero.

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classification

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der Maarel,

in the 1980s. In:

science
E., 1984. Vegetation
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re
(eds), Trends
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Plenum
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explicativas

1963. Los bosques

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27: 454-476.

1972. Ecological

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entre

clim?ticas

variables

of cluster

66: 846-850.

Assoc.

Statist.

1968. Correlaci?n
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for the evaluation

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1971. Objective

G.,

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Van

hypotheses

14th Congr.

1941. Exploraciones
1958.

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Soc.

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Nat.

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de

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1956. Studies

L.,

F.,

UNESCO,

1937.

H.,

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formaciones

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las

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con breve

F.,

ac
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Bolet?n
Informativo

de la Rep?bli
Tamayo, F., 1964. Mapa
fitogeogr?fico
preliminar
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Ministerio
de Agricultura
y Cr?a, Caracas.

Caracas.
H.,

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Paraguan?.

Tamayo,

de

Renova

Austin.

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3, FUDECO,

Tamayo,

y desarrollo

Naturales

de Recursos

y Cr?a, Direcci?n

Agricultura

of Texas

fitogeogr?fico
Ministerio

forestal.

agropecuaria

of arid vegetation
in tropical
of desert biota.
(ed.), Evolution

In: D. W. Goodall

Smith, R. F., Ferrer V. E. & Chavez, A.,

tual de la regi?n
centro-occidental.
No.

1985. Multiple
purposes
9:
inventory. Environ. Manage.
L.,

1976. Evolution

G.,

America.

9: 217-224.

Conserv.

A. & Pia,

S. D., Colma,

Matteucci,

& Pia,

Environ.

State.

of Falc?n

maps

A.

Colma,

Sarmiento,

subtropical
Accepted

9.10.1986.

I2?00'

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S. MATTEUCCI,THE VEGETATIONOF FALC?N STATE, VENEZUELA. VEGETATIO 70: 67-91,1987

FALCON STATE,VENEZUELA
PHYSIOGRAPHIC PROVINCES

cJiL8e^N

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I Io00

FIELD DATACOLLECTEDFROM 1976 TO

1978

BASEMAPCOMPILED
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SUPPLIED BY CARTOGRAF?A

CLOUD FOREST
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