Landscape and Urban Planning 77 (2006) 240–254

Unplanned land clearing of Colombian rainforests:

Spreading like disease?
Andres Etter a,b,c,∗ , Clive McAlpine a,b , Stuart Phinn a,b ,
David Pullar a,b , Hugh Possingham a
a The Ecology Centre, The University of Queensland, Brisbane, Qld 4072, Australia
b Centre for Remote Sensing and Spatial Analysis, School of Geography, Planning and Architecture,
The University of Queensland, Brisbane, Qld 4072, Australia
c Facultad de Estudios Ambientales y Rurales, Universidad Javeriana, Bogotá, Colombia

Received 5 December 2004; received in revised form 1 March 2005; accepted 2 March 2005
Available online 25 May 2005

Abstract

Deforestation often occurs as temporal waves and in localized fronts termed ‘deforestation hotspots’ driven by economic pulses
and population pressure. Of particular concern for conservation planning are ‘biodiversity hotspots’ where high concentrations
of endemic species undergo rapid loss and fragmentation of habitat. We investigate the deforestation process in Caquetá, a
biodiversity hotspot and major colonization front of the Colombian Amazon using multi-temporal satellite imagery of the periods
1989–1996–1999–2002. The probabilities of deforestation and regeneration were modeled against soil fertility, accessibility and
neighborhood terms, using logistic regression analysis. Deforestation and regeneration patterns and rates were highly variable
across the colonization front. The regional average annual deforestation rate was 2.6%, but varied locally between −1.8%
(regeneration) and 5.3%, with maximum rates in landscapes with 40–60% forest cover and highest edge densities, showing an
analogous pattern to the spread of disease. Soil fertility and forest and secondary vegetation neighbors showed positive and
significant relationships with the probability of deforestation. For forest regeneration, soil fertility had a significant negative
effect while the other parameters were marginally significant. The logistic regression models across all periods showed a high
level of discrimination power for both deforestation and forest regeneration, with ROC values >0.80. We document the effect
of policies and institutional changes on the land clearing process, such as the failed peace process between government and
guerillas in 1999–2002, which redirected the spread of deforestation and increased forest regeneration. The implications for
conservation in biologically rich areas, such as Caquetá are discussed.
© 2005 Elsevier B.V. All rights reserved.

Keywords: Tropical deforestation; Colonization; Multiple scales; Biodiversity hotspots; Colombia

1. Introduction
∗ Corresponding author. Tel.: +61 7 33653535;
fax: +61 7 33656899. The clearing of forests by humans has been go-
E-mail address: andres.etter@uq.edu.au (A. Etter). ing on for at least 10,000 years since the advent of
0169-2046/$ – see front matter © 2005 Elsevier B.V. All rights reserved.
doi:10.1016/j.landurbplan.2005.03.002
 A. Etter et al. / Landscape and Urban Planning 77 (2006) 240–254
agriculture and ancient civilizations, but rapidly in-
creased during the late 1800s and 1900s with the ex-
pansion of the human population and the industrial
economy. Since 1900, the areas of cropland have dou-
bled (Houghton, 1994), with the rate of deforestation
accelerating significantly since the 1960s, coinciding
with rapid global population growth, especially, in
the tropics. Land use change often occurs in tempo-
ral waves and in localized fronts termed ‘deforestation
hotspots’ by Myers (1993), that respond to the pulses
of change in land use drivers (e.g. Lambin and Ehrlich,
1997; Sierra, 2000; Viña et al., 2004). Of high concern
for conservation planning are ‘biodiversity hotspots’
with high concentrations of endemic species that are
undergoing exceptional loss of habitat (Myers et al.,
2000). These hotspots are mostly located in the trop-
ical regions, and often coincide with active coloniza-
tion fronts. The Amazonian foothills in the Caquetá and
Putumayo Departments of Colombia are renowned for
their high species richness and high levels of endemism
(Hernández et al., 1992). These areas correspond to one
of the hypothesized Amazonian Pleistocene Refugia
“Napo-Putumayo”. However, most of the eastern Ama-
zon region of Colombia has been extensively cleared
(Etter, 1998) with only one conservation area (La Paya
National Park) to the south in the border with Peru.
By definition, landscapes are dynamic, with change
an inherent characteristic of the interaction between
nature and society (Bürgi et al., 2004; Forman, 1995).
Understanding and predicting landscape dynamics is
an expanding research field in landscape ecology (Wu
and Hobbs, 2002). One of the most obvious changes in
landscape structure is change in land cover due to clear-
ing. Land cover changes occur at multiple spatial and
temporal scales and in varying biophysical and socio-
economic contexts (Wiens, 2002); thereby limiting our
ability to generalize about this process. In tropical re-
gions, land clearing has been studied mainly at regional
(10,000 s km2 ) or national scales (100,000 s km2 ) (e.g.
Kleinn et al., 2002; Laurance et al., 2001; Skole et al.,
1994), often overshadowing the complexity and vari-
ability at finer scales. Local scale (100–1000 s km2 )
studies, however, are not always representative of the
broader regional and national contexts. Understanding
how the different scales interact is a prerequisite for
a holistic understanding of the patterns and processes
of landscape change. The difficulties of mapping and
monitoring land cover change have been largely over-
241
come with the expansion of earth observation satel-
lites technology, which provide greatly increased ac-
curacy and precision. However, the lack of appropriate
research methods and comparable datasets has led to
contradictory figures on the rates and patterns of de-
forestation, even in small countries with accurate land
cover data, such as Costa Rica (Kleinn et al., 2002).
In the tropics, unlike most temperate regions, land
clearing and especially, deforestation, is still the dom-
inant land cover change process (Lucas et al., 2004).
Deforestation occurs due to a wide range of economic,
political and demographic reasons, which need to be
adequately addressed if their impacts are to be un-
derstood, and the resulting knowledge applied in pol-
icy and management. For example, deforestation has
been often actively promoted by governments through
constructing infrastructure, such as roads (Wilson
et al., 2004), and allocating parcels of land to migrat-
ing colonists in a semi-planned way. This is the case in
the Brazilian Amazon regions, such as Rôndonia (Dale
et al., 1993), and also Ecuador (Sierra, 2000; Viña et al.,
2004). In other cases, land clearing is a more sponta-
neous and uncontrolled process, as occurs in coloniza-
tion fronts of Colombia and Central America (Etter and
Andrade, 1987; Turner et al., 2001; Viña et al., 2004).
There also is community controlled deforestation, such
as used by indigenous communities in the Amazon,
where the allocation of subsistence cropping areas is
done according to social rules (e.g. van der Hammen,
1992).
Not only change, but also persistence of land cover,
is an important aspect when studying land cover
dynamics (Bürgi et al., 2004; Pontius et al., 2004).
Persistence of forests in the landscape requires consid-
ering that deforestation can result in both, permanent
transformation (e.g. cattle ranching in most tropical
lowlands of Latin America), or transitory change (e.g.
the traditional migratory “slash-and-burn” agriculture
throughout most of the tropics). Under the latter
traditional system, fallow systems were extensively
used and forest regeneration occurred. However, due
to increased population pressure, this practice has
become less common and the length of fallow periods
reduced, often preventing effective forest regeneration.
In Colombia until the early 1900s, most of the
economic activity and land clearing was located in the
Andean region and the Caribbean lowlands. During the
1900s, colonization steadily expanded into the humid
242 A. Etter et al. / Landscape and Urban Planning 77 (2006) 240–254
lowlands (Palacios, 2001), which were largely depop-
ulated from European settlers, but were still home to
many low-density indigenous communities. Between
1830 and 1930, over 2.5 million hectares of so-called
“baldios” (empty lands) owned by the State were
progressively handed over to individuals (Legrand,
1988). After 1950, a more spontaneous, uncontrolled
colonization of the State owned lands occurred, mainly
in the forested lowlands of the Amazon, Orinoco and
middle Magdalena regions. From the 1970s, the rate of
deforestation increased significantly due to migration
from the densely populated Andean region. The latest
national estimates of deforestation compiled by the
Ministry of Environment are 100,000 ha/year (Instituto
de Estudios Ambientales, IDEAM, 2004), however,
the data and calculations employed to generate this
figure are unknown.
Colonization in the eastern Andean foothills be-
tween 1940 and 1970 are described by Brücher (1974).
Until the 1930s, the main economic activities in this
region were the harvesting of quinine and rubber. In
1935, a large cattle property “Larandia” was cut into
the rainforests east of Florencia, after which a few
colonists followed, which marked the onset of mod-
ern land clearing in the region and the commencement
of a major active colonization front. In 1959, through
a concession to the Agrarian Bank of 700,000 ha, a
government sponsored colonization scheme began in
Fig. 1. Location of study area showing: (a) Caquetá region in the country and (b) location of the five 100 km2 sample areas used for landscape-level
analysis.
three areas within the Caquetá region in the Amazon
lowlands. From 1961, the Colombian Institute of Land
Reform continued this scheme. The approach consisted
of delimiting six areas of about 10,000 ha, within which
land parcels, 50–100 ha in size were allocated, and im-
migrating colonists could access state funded credits
and legal titles. Unlike the colonization schemes of
Brazil in the 1970s, the Colombian schemes of the early
1960s had less infrastructure development and ended
in waves of uncontrolled colonization. From the late
1980s, three main forces have driven the colonization
process in Colombia: landlessness, illicit crops, and the
presence of rebel armies.
This paper describes and models the deforestation
process in the Caquetá Department in the Amazon re-
gion of Colombia (Fig. 1) from 1989 to 2002. We have
three specific objectives. First, to map and quantify
rates and patterns of deforestation and forest regenera-
tion at different spatial scales and locations within the
Caquetá colonization front. Second, to analyze the re-
lationship between deforestation/regeneration and the
changing demographic and political context of the re-
gion. Third, to model the relationship between forest
cover change and biophysical and socio-economic fac-
tors. In conclusion, we discuss how the findings could
be applied in conservation planning for biologically
rich but threatened landscapes areas that are biodiver-
sity hotspots.
 A. Etter et al. / Landscape and Urban Planning 77 (2006) 240–254
2. Data and methods
2.1. Study area
The study area (23,000 km2 ) is located in the Ca-
quetá Department in the Andean foothills of the eastern
Amazon region of Colombia (Fig. 1a). It includes 17
municipalities (whole or in part) with a total rural pop-
ulation of about 180,000 in 1993. Population densities
range from 7 to 16 inhabitants/km2 of cleared land.
The most extensive land use is cattle ranching, with
illegal crops of coca (Erythroxylum coca) covering a
smaller area but increasing in economic importance
since the early 1980s. Latest figures indicate a drop
of almost 50% in illegal plantations area in Colombia
from a high in 2001 (United Nations Office for Drug
Control, UNDOC, 2004).
Caquetá Department comprises mainly flat to rolling
plains extending east of the eastern Andean Range, with
an average altitude of 200 m (Carvajal et al., 1993). The
climate is humid tropical with an average annual rain-
fall ranging from 3500 mm in the west to 2500 mm
in the east, and with 1–2 dry months with less than
100 mm. Soils are very acid and deep, except for some
recent alluvial deposits, and mostly well-drained ex-
cept for flat river terraces (Malagón et al., 1993). The
natural vegetation is composed of several rainforest
types varying in structure and composition, depend-
ing on physical conditions, notably soil, relief and
drainage. Standing biomass varies between 210 and
280 t/ha. The forests of the well-drained interfluves
are 25 m high, and have higher species counts than
the alluvial forests. Carvajal et al. (1993) recorded
145 species of trees >10 cm DBH (diameter at breast
height) occurring in forests with different levels of dis-
turbance and fragmentation in the region. A decrease
in species numbers and an increase in abundance of
palms was observed in fragments and disturbed forests.
Common trees included species of the genus Eschweil-
era, Brosimum, Ocotea, Virola, Pouteria, Parkia and
Clathrotropis, and palms of Jessenia, Iriartea and Mau-
ritia.
2.2. Data preparation
Four Landsat 5-Thematic Mapper (TM) and Land-
sat 7-Enhanced Thematic Mapper Plus (ETM+) images
from 1986, 1996, 1999 and 2002 were used to extract
243
land cover maps (images downloaded from the Global
Land Cover Facility of the University of Maryland,
http://glcfapp.umiacs.umd.edu:8080/esdi/index.jsp).
The exact period between images was calculated using
the acquisition dates, to 6.65 years for the 1989–1996
period, 3.25 years for the 1996–1999 period and 2.15
years for the 1999–2002 period. All images were
geo-referenced to the Landsat 7 (ETM+) 2002 image
to a level of less than 0.3 root mean standard error
(R.M.S.E.), or 9.0 m. Radiometric normalization was
not applied as each image was classified independently
and the land cover types mapped exhibited signif-
icantly different spectral signatures. A supervised
classification procedure using the ERDAS-Imagine
software was applied to differentiate 11 classes:
non-flooded forest; flood forest; non-forest secondary
vegetation; flooded scrublands; crops; pastures;
savannas; burnt areas; water bodies; cloud-shadows.
Both forest classes were merged into one “forest”
class. Due to difficulties in consistently differentiating
pastures, savannas, crops, fire and barren land, they
were merged into a “cleared” class. The images from
1996 and 1999 showed a partial cover of clouds, which
was overcome through reclassification by comparison
of the earlier and later dates. The final classification
included four classes: forests, non-forest secondary
vegetation, cleared and water. Due to lack of suitable
reference data for the different image dates, a pseudo
error assessment was conducted by classifying the
training sites using the classification algorithm of the
ERDAS-Imagine software module, and the accuracy
of the classification reported (Jensen, 1996). Although
this is a biased approach to error assessment, it does
indicate the suitability of training sites for each land
cover class of each image. The accuracy of the training
sites used was assessed using contingency tables,
which showed agreement levels of more than 95% for
all classes in all time-steps.
To prepare the images for change analysis, the
image-to-image geometric registration accuracy of
the classified images was assessed using the method
outlined by Phinn and Rowland (2001). This approach
directly compared the coordinates of 25 well-identified
pixels, or ground control points in each pair of images
1989–1996, 1996–1999 and 1999–2002. The extent
of spatial misregistration between images showed a
maximum value of 0.29 (±0.16 S.D.) pixel or 8.7 m
for 1989–1996, 0.17 (±0.0.09 S.D.) pixel or 5.1 m
244 A. Etter et al. / Landscape and Urban Planning 77 (2006) 240–254
for 1996–1999, and 0.16 (±0.11 S.D.) pixel or 4.8 m
for the 1999–2002 period, all below the maximum
acceptable value of 0.5 pixel (Jensen, 1996).
In extensively cleared areas, the classified im-
ages contained numerous isolated individual pixels
(<0.1 ha) classified as “secondary vegetation” or “for-
est”. These isolated pixels are smaller than remnant
forest patches, and were treated as noise (one such
pixel could correspond to one tree in a sea of grass
and would not be considered “secondary vegetation” or
“forest”). To decrease this high spatial frequency noise
or image speckle, a 3 × 3 Majority Filter (equivalent to
ca. 1 ha, which subsequently became the unit of analy-
sis) was applied to all images. Although this procedure
smoothed the land cover map by reducing some spatial
detail, it sharpened the spatial patterns of forest and
secondary vegetation. An experimental test with 1996
data indicated that by using a 3 × 3 matrix only 0.4%
of the area was lost, while for other land cover aspects,
such as edge density, where relative values were similar
although absolute edge density decreased. The images
were then re-sampled to a uniform 1 ha grid, in order
to perform the land cover change detection procedure.
For each image date, a forest–non-forest (1, 0) map
was created. Transition maps for forest–non-forest (de-
forestation) and non-forest–forest (forest regeneration)
were produced for the 1989–1996, 1996–1999 and
1999–2002 intervals. Because of the short time inter-
vals between land cover maps, we assume that regen-
eration only occurs when there is a transition from
secondary vegetation to forest, and not directly from
cleared land or water. In studies, such as Nagendra et al.
(2003), time intervals between dates are too short (<10
years) for succession to regenerate a forest cover effec-
tively from cleared. To test the accuracy of regeneration
transitions, we checked for the proportion of direct tran-
sitions from cleared or water to forest, which occurred
for the 1989–1996 in 0.03% of cases. For 1996–1999
and 1999–2002, no “false transitions” were recorded.
The “false transitions” of the 1989–1996 period were
treated as errors and eliminated from the forest regen-
eration map. The lack of false transitions was further
confirmation of the accuracy of the maps.
A neighborhood map of forest and secondary
vegetation was calculated for each forest–non-forest
map using a 5 × 5 moving window (25 ha). In order to
evaluate the effects of physical and economic factors
on the spatial distribution of the forest mosaic, data
on soils and accessibility were retrieved from existing
spatial data. Soil fertility maps with three classes
(1–3) treated as continuous were derived from existing
general soil maps from the National Geographic In-
stitute (Instituto Geografico Agustin Codazzi, IGAC,
1983). To take into account the accessibility of a site, a
cost–distance map was produced for each time-step as
a measure of physical and economic accessibility. This
was done using spatial data on rivers, road infrastruc-
ture, terrain/topography and surrounding settlements
derived from remote sensing images and topographic
maps (scale 1:100,000) sourced from the Instituto
Geografico Agustin Codazzi (IGAC). Differential
friction values were then assigned using Arc View 3.2
GIS, with the values proportional to the travel time
starting with paved roads (1), unsealed (5), tracks
(10), major rivers (3), minor rivers (15) and plain land
(25).
Five 100 km2 subsets were chosen along the col-
onization front taking into account the regional colo-
nization age axis gradient (west–east), to evaluate the
differences in landscape pattern and rates of change,
thereby providing a landscape-level comparison with
the regional analyses (Fig. 1b). The location of the sub-
sets was chosen in order to be: (a) spread over the entire
colonization front and (b) covering the entire gradient
of remnant forest proportion (<10 to >90%).
2.3. Logistic regression models
Logistic regression models, a variation of ordinary
regression when the dependent variable is binary (0,
1), were applied to the binary 1989–1996, 1996–1999
and 1999–2002 transition maps of deforestation and
forest regeneration to determine the effect and relative
importance of four explanatory variables: soil fertil-
ity, cost–distance and land cover neighborhood (for-
est and non-forest secondary vegetation). Modeling
was performed using R public-domain statistical pack-
age (R Project for Statistical Computing Release 1.9.0
http://www.r-project.org). Explanatory variables used
in the model were standardized using the SCALE func-
tion in R.
log it(y) = β0 + β1 X1 + · · · + βk+1 Xk+1 (1)
For the analysis in R, a random sample of 20,000
cells (1%) was extracted from each transition map
because of the extremely large number of cells
 A. Etter et al. / Landscape and Urban Planning 77 (2006) 240–254
(>2,000,000), and also to avoid problems associated
with spatial autocorrelation in the explanatory vari-
ables.
Second, in order to test for the consistency of the
models through time and the possibility to explain
subsequent clearing events, a logistic regression using
the same set of explanatory variables, was also applied
to the full maps using Idrisi GIS function “logisti-
creg” (Clark-Labs, 2001). Models developed for the
1989–1996 period were independently validated for
the 1996–1999 period, and the 1996–1999 models
validated for the 1999–2002 period. These analyses
quantified temporal similarities in the spatial patterns
of deforestation and forest regeneration for the region.
The Idrisi function validates the discrimination ability
of the model by using the area under the receiver
operator characteristics or ROC curve (Metz, 1978).
The application of the ROC to predicting land cover
changes is discussed in detail in Pontius and Batchu
(2003). It measures the agreement of the predicted
and the observed values of change, in terms of the true
positives (correct change) against the false positives
(errors) in the predicted model. The better the model
discriminates the observed data, the more likely it is
that the cells with a high predicted probability at t1 (e.g.
1989–1996) will have changed (deforestation or re-
generation) in the observed data at t2 (e.g. 1996–1999).
A random prediction will have a ROC close to 0.5,
while a perfect prediction would have a ROC of 1.
The importance of the ROC method is that it analyses
independently the specification of location from the
specification of quantity (Pontius and Batchu, 2003).
2.4. Rates of change
The annual rates of change for the three land
cover classes (forest, secondary vegetation and cleared
land) were calculated for the whole region and for
the 100 km2 windows, using the formula (Puyravaud,
2003):



1 Ai2
rate = ln
t2 − t 1 Ai1
where Ai1 is the cover of class i at an initial time (t1 )
and Ai2 is the cover of class i at a later time (t2 ). The
deforestation rates were also calculated by Municipal-
ity, and then regressed against rural population growth
data, derived from the 1985 and 1993 national census
245
data (Departamento Administrativo Nacional de Es-
tadistica, DANE, 1993).
3. Results
3.1. Regional patterns of land cover change
By 1986, the study area was already significantly
cleared (52%), especially, in the west of Caquetá
Department near Florencia. However, the region
continued to experience steady deforestation during
the subsequent 13 years, with 66% cleared by 2002
(Fig. 2). During this period, some 320,000 ha of
forest were cleared at an average annual rate of 2.4%
(approximately 25,000 ha/year). There were, however,
significant variations between the analyzed periods;
especially, noteworthy is the substantial drop in defor-
estation in the 1999–2002 period. The deforestation
rates peaked at more than 40,000 ha/year or 4.1%
during 1996–1999, while total land clearing reached
66,660 ha/year or 7.8% when secondary vegetation
(40% of all clearing) was also included.
An important compensating relationship between
deforestation and forest regeneration was found during
the land clearing process, where deforestation was
partially offset by parallel regeneration of forests in
previously cleared land (Fig. 3). The effect of forest
regeneration on net forest change was evident for
all three periods, with the highest effect during
1999–2002. From 1989 to 2002, forest regeneration,
on average, counterbalanced deforestation by 27.5%,
with the lowest value of 17.7% for the 1996–1999
period. The spatial pattern of deforestation was
concentrated along an active colonization front, while
Fig. 2. Regional values of area of the three main land cover classes
for the four mapping dates.
246 A. Etter et al. / Landscape and Urban Planning 77 (2006) 240–254
Fig. 3. Net forest cover change as a result of the effect of forest
regeneration on deforestation during the land clearing process for
the 1989–2002 period.
forest regeneration was found more in the highly
cleared areas of the region (Fig. 4).
Analysis of the individual annual rates of transition
between the three land covers (forest, secondary veg-
Fig. 4. Time series of maps of forest cover change showing: (a) forest cover for all dates in black and for the transition periods in black; (b)
deforestation; (c) forest regeneration.
etation, cleared land) for the three periods as captured
by each Landsat TM/ETM + image pair provides fur-
ther detail on spatial and temporal variations in defor-
estation (Fig. 5). The most dynamic component of the
landscape is secondary vegetation with gross annual
rates of change up to 14.5% (1996–1999 period), in-
cluding either re-clearing (12.4%) or more permanent
forest regeneration (2.1%). Forests were transformed at
total rates of between 2.8 and 4.1%. Part of the forests
went to cleared land (1.4–2.1% annual rates), while a
similar amount reverted to non-forest secondary vege-
tation (1.4–1.9%). The latter would mean that part of
the cleared forests undergo very rapid abandonment in
some areas, which are then partially re-cleared before
reaching a forest state.
The change matrices of each period for the entire
region show that the probability of persistence of
forest for the 1989–1996, 1996–1999 and 1999–2002
 A. Etter et al. / Landscape and Urban Planning 77 (2006) 240–254 247

Fig. 5. Annual rates of change of land cover transitions for forest, secondary vegetation and cleared land, for the three studied periods. Size
of boxes is equivalent to the relative proportion of cover in the landscape. Thickness of arrows indicate the relative intensity of change, and
numbers are relative to the amount of cover type.

transition periods increased from 0.81 to 0.84 and 0.88, 3.2. Changes at the landscape level
respectively (Table 1). Stability of the cleared area
similarly increased from 0.82 to 0.97 for 1996–1999 The annual rates of change calculated for the entire
and decreased slightly to 0.91 for 1999–2002. Sec- region (2.4%) mask the landscape-level heterogeneity
ondary vegetation was a more dynamic landscape of spatial and temporal patterns in the spread of colo-
component, with only 0.45–0.66 persistence of its nization. Fig. 6a provides an example of three 100 km2
cells during the transition periods. The stability of the sample areas corresponding to different phases in the
water cells decreased as a result of increasing channel colonization process, showing that forests disappear
migration. However, at the local level, the persistence and connectivity is lost much faster in the middle phase
of the three main land covers varied greatly, depending (ii) than in phase (i) and the later phase (iii). Although
on the proportion of forest cleared. the proportions of forest and cleared land are differ-
ent, it is notorious that the general spatial patterns of
Table 1
transformation are strikingly similar, fitting into a trans-
Transition matrices of land covers for the 1989–1996, 1996–1999 formation gradient, of decreasing forest and increasing
and 1999–2002 periods for the Caquetá colonization front cleared land, and indicating a temporal continuity in
Forest Secondary Cleared Water the eastward progression of the front. Consequently,
vegetation areas in the east exhibit similar patterns to the ar-
From 1989 to 1996 eas in the west of the region, which underwent clear-
Forest 0.81 0.09 0.09 0.00 ing several years ago. By coupling the trajectory
Secondary vegetation 0.15 0.45 0.38 0.00 1989i–1996i–1999i–2002i–1989ii–1996ii–1999ii, . . .,
Cleared 0.04 0.12 0.82 0.01 –2002iii, we can predict that the transformation cy-
Water 0.00 0.01 0.06 0.89
cle from forest to cleared at the landscape level in the
From 1996 to 1999 region takes approximately 40 years. This is further
Forest 0.84 0.07 0.08 0.00
demonstrated by Fig. 6b, where the rates of change for
Secondary vegetation 0.08 0.45 0.47 0.00
Cleared 0.00 0.03 0.97 0.00 the three periods of all the five 100 km sample areas
Water 0.00 0.00 0.02 0.77 (n = 15) are plotted against remnant forest proportions.
From 1999 to 2002
The rate of change curve has a quadratic shape with the
Forest 0.88 0.05 0.07 0.00 highest annual rates of about 5% found in areas with
Secondary vegetation 0.12 0.66 0.23 0.00 forest proportions between 0.3 and 0.7. However, be-
Cleared 0.00 0.09 0.91 0.00 low 0.3 and above 0.7 forest cover, the change rates fall
Water 0.01 0.01 0.27 0.71 significantly below 2.5%. Therefore, the regional aver-
248 A. Etter et al. / Landscape and Urban Planning 77 (2006) 240–254

Fig. 6. Land cover change at the local scale in three 100 km2 sample areas: (a) maps of land cover (forest, black; secondary vegetation, gray;
cleared, white) corresponding to: (i) early stage (window 4, in Fig. 1); (ii) intermediate stage (window 2); (iii) late stage (window 1) of the
colonization process; (b) relation between rates of change and forest proportion at the local scale; (c) relation between rates of change and forest
edge density at the local scale.

age of 2.4% obscures these important local-scale dif- ity level. The annual average for Caquetá Department
ferences, with the spread of transformation varying by was 0.13 ha/inhabitant, with a maximum of 1.07 in the
several orders of magnitude across a colonization front. east (La Macarena), and a minimum of −0.128 (forest
Deforestation was significantly higher in land- regeneration) in the west (Milan). This indicates that,
scapes with highest edge density (Fig. 6c), indicating in the active colonization front, an average family of
that the rate of change is correlated with exposed forest
edge. The effect of population growth rates at the
municipality level on the rate of forest clearing (Fig. 7)
shows a positive relationship, with high deforestation
rates occurring in municipalities with higher rural
population increase. Correspondingly, municipalities
with a net increase in forest area (positive forest
regeneration) correspond to municipalities, mainly
located in the western part of Caquetá Department,
with a net emigration of rural population. Therefore,
the spatial pattern of high rates of deforestation
along the colonization front moves from west to east,
tracking parallel changes of rural population densities. Fig. 7. Relationship between rural population growth rate of mu-
Table 2 shows the cleared area per rural inhabitant nicipalities (1985–1993) and forest clearing rate (1989–1996) in the
per year for the 1989–1996 period at the municipal- Caquetá region.
 A. Etter et al. / Landscape and Urban Planning 77 (2006) 240–254 249

Table 2
Municipality aggregated data of, annual forest clearing per rural inhabitant, rural population growth rates, and difference with national rural
population growth rate, in the Caquetá colonization front during 1989–1996
Municipality Annual deforestation per rural Annual (%) rural population % Difference to national population
inhabitant (ha) (1989–1996) growth rate (1985–1993) growth rate (1985–1993)
La Macarenaa 1.07 0.092 485
C. del Chaira 1.04 0.165 954
Solanoa 0.51 0.084 434
San Vicente 0.28 0.055 248
Valparaı́soa 0.10 0.203 1196
La Montanita 0.06 0.032 103
Albaniaa −0.01 −0.039 −347
B. de los Andaquı́es −0.01 −0.044 −379
Florenciaa −0.01 −0.037 −334
Curilloa −0.01 0.010 −35
Puerto Rico −0.01 −0.023 −244
El Paujil −0.02 0.040 157
EL Doncello −0.03 −0.015 −196
Morelia −0.04 0.024 50
Milan −0.03 0.117 649
Average 0.13 0.047 192
a Municipalities with more than 30% of their area out of study area.

five persons cleared between 1 and 5 ha/year during
this period. Also, municipalities that experienced ac-
tive deforestation have a rural population growth rate
up to 10 times the national average, while the opposite
trend is true for municipalities, where regeneration of
forests is predominant (Table 2).
3.3. Predictions of deforestation and forest
regeneration
An objective of this study was to investigate the ef-
fect and relative importance of explanatory variables,
such as soil fertility, accessibility and forest neighbors
on forest transitions. The results of the logistic regres-
sion models show that the explanatory variables have
a stronger influence on deforestation than forest regen-
eration (Table 3).
Soil fertility shows a consistent opposite effect
for both processes, with a positive effect for defor-
estation and negative effect for forest regeneration.
The significant parameters for cost–distance have a
negative effect for deforestation and positive effect for
regeneration. Forest and secondary vegetation neigh-
bors showed a positive and significant relationship

Table 3
Standardized parameter estimates of explanatory variables derived from the logistic regression analyses: (a) deforestation and (b) forest
regeneration
Period Intercept Soil Cost–distance Forest neighbors Secondary vegetation neighbors
Part (a)
1989–1996 −3.08∗ 0.39∗ −0.25∗ 1.32∗ 0.89∗
1996–1999 −3.18∗ 0.06∗ −0.18 0.72∗ 0.82∗
1999–2002 −3.64∗ 0.03 −0.08 0.65∗ 0.67∗
Part (b)
1989–1996 −15.23 −0.55∗ −0.12 −0.02 −33.79
1996–1999 4.49∗ −0.24∗ −0.10 0.54∗ 0.48∗
1999–2002 14.24 −0.43∗ 0.33∗ 0.05 31.01
Regression is from random samples (n = 20,000).
* Significant at p < 0.01.
250 A. Etter et al. / Landscape and Urban Planning 77 (2006) 240–254
Table 4
Receiver operating characteristics (ROC) values of the predictive
model of one date applied to observed values of subsequent dates:
(a) deforestation and (b) forest regeneration
Predicted Observed
1996–1999 1999–2002
Part (a)
1989–1996 0.785 0.808
1996–1999 – 0.824
Part (b)
1989–1996 0.802 0.803
1996–1999 – 0.841
with deforestation for all transition periods. For forest
regeneration, however, these variables were only sig-
nificant for the 1996–1999 with a positive effect. The
relative importance of the scaled explanatory variables
shows a predominance of the neighborhood variables
of forest and secondary vegetation, with parameter
values for the neighborhood effects two to three times
higher than for soil fertility and cost–distance.
The predictive ability of the models generated using
Idrisi for the successive periods was surprisingly high
(Table 4). The area under the ROC curve for defor-
estation and regeneration for all change periods ranged
from 0.785 to 0.841, indicating a moderate to high level
of discrimination, and pointing to spatial patterns of
transformation being fairly constant along the trans-
formation process across the region.
4. Discussion and conclusions
We have shown that (i) patterns and rates of change
vary significantly within a spontaneous colonization
front; (ii) the rates of deforestation and forest re-
generation are closely related to the spatial patterns
of forest cover; (iii) the rates of change of the rural
population are strong covariates with deforestation and
regeneration. Drawing on an epidemiological analogy,
we infer that the land clearing process mimics the
spread of disease (Medlock and Kot, 2003), following
a negative sigmoid pattern, with rates peaking at
intermediate forest cover values, where the amount of
exposed forest edge is at its maximum, and therefore,
the “infection probability” of forests to clearing
maximizes.
The driving forces of land cover change, especially,
deforestation, result from the complex interaction of
socio-political and economic processes. The major
drivers of deforestation in the eastern Amazon region of
Colombia during the last 20 years have been the illegal
economy of coca crops and cattle ranching. Our study,
which extended from 1989 to 2002, shows the highest
deforestation rates occurred between 1996 and 1999,
when the illegal economy of narcotics was booming
in the region (United Nations Office for Drug Control,
UNDOC, 2004). During this period, the region was a
stronghold of the FARC guerrillas who controlled the
political and economic activity over large areas. It has
been debated what effects the Colombian armed con-
flict has on the deforestation processes (Dávalos, 2001).
Our results show an increased deforestation during the
1996–1999 period, had a positive link with high guerri-
lla activity and low government presence. The presence
of guerilla armies poses a major obstacle to managing
deforestation in a planned manner, and prohibits any
form of conservation planning and management.
4.1. Regional and landscape-level land cover
changes
The average annual rate of clearing for Caquetá dur-
ing 1989–2002 was 25,000 ha, with a peak of 41,000 ha
during 1996–1999. This indicates that annual national
rate of 100,000 ha reported by Instituto de Estudios
Ambientales (2004) is probably an underestimate by
25–40%, as the Caquetá colonization front is one of
several similar deforestation fronts in Colombia (e.g.
Putumayo, Guaviare–Vichada, Middle Magdalena, Pa-
cific lowlands). The average rates of deforestation
across the region broadly match the results provided
by other studies (Sanchez-Azofeifa et al., 2001; Sierra,
2000; Viña et al., 2004). Sierra (2000) showed that rates
of change across a colonization front in the Ecuado-
rian Amazon region are highly variable. However, our
study provided a more spatially explicit assessment of
the rates and spatial variation of deforestation by inves-
tigating the relationship between the rates of change,
clearing phase and spatial pattern of forest cover. This
analysis indicated that peak of deforestation occurs
when the remnant forest cover is around 50%, and the
exposed forest edge is at its maximum.
From a spatial pattern perspective, the probability
of deforestation at the landscape-level is related to the
 A. Etter et al. / Landscape and Urban Planning 77 (2006) 240–254
proportion of remnant forest (Lambin and Ehrlich,
1997), and to the proximity and amount of forest edge
(Ludeke et al., 1990). Highest rates of deforestation
are expected in areas with intermediate forest cover
(35–80%) and highest edge density. Our results
confirm this relationship for the Colombian Amazon
in the 1989–2002 period. Edges are symptomatic
of clearing, but also facilitate clearing by exposing
the remaining forest to settlers armed with axes and
chainsaws. This result suggests a correlative, but also a
partly causal relationship between landscape structure
(forest proportion and edge density) and the rate of
clearing of Colombia’s Amazonian forests. A similar
strong ‘spread effect’ of deforestation was observed
in Honduras by Ludeke et al. (1990), as forest clearing
tends to spread from forest edges or from previous
forest openings. In a fragmented forest, more edge
per unit area is exposed to human activity, resulting in
increased human accessibility to the interior of forests.
Forest fragmentation and increased edges are also
symptomatic of human settlements. Such settlements
are likely to become centers of diffusion of deforesta-
tion activities if there is a demographic, economic, or
social process driving an increase in forest clearance.
Deforestation is not a terminal event, but rather
a setting in motion of complex forest clearing and
regeneration processes (Nagendra et al., 2003; Nepstad
et al., 1991). Our results show that these counter-
balancing processes have definite spatial patterns.
The highest rate of regeneration occurs in the more
transformed areas, less than 80 km from Florencia,
the main economic center of the region, while
increased deforestation occurs at distances greater
than 80–100 km. The deforestation front exhibits a
quadratic function, with the rate of clearing peaking
in the 80–140 km distance zone, slowly shifting to the
east as colonization proceeds. This is consistent with
the pattern described by Mertens and Lambin (1999)
for Cameroon where maximum deforestation was
found at intermediate distances, indicating a general
pattern in spontaneous colonization fronts.
Socio-economic and political variables are impor-
tant drivers of land use change and/or persistence
(Bürgi et al., 2004; Lambin et al., 2001). For example,
deforestation rates and patterns vary across national
borders in response to different political and economic
contexts, such as government planned (Ecuador) and
unplanned (Colombia) colonization processes (Viña et
251
al., 2004). Caquetá went through significant political
and socio-economic changes due to the peace process,
which the government of President Pastrana agreed
to with the irregular leftist armed forces FARC, from
December 1998 to March 2002. This meant the with-
drawal of the military from 42,000 km2 including two
municipalities of our study area (La Macarena and San
Vicente del Caguán). The significant drop in the defor-
estation rates between 1999 and 2002 can be primarily
attributed to the political and economic conditions that
prevailed in the military free zone during this period.
If the process was effective, a decline of both the le-
gal and illegal economic activities would be expected
in these municipalities (La Macarena and San Vicente
del Caguán) during the peace process. However, these
two municipalities contributed to almost 80% of the
regional clearing in 1999–2002, compared to between
30 and 50% prior to 1999. This indicates that the il-
legal economy was still largely active in the exclusion
zone, giving support to the claims of the government
that the rebels were using the area for illegal economic
activities during the peace process. In contrast over the
Caquetá Department for the same period, deforesta-
tion slowed down while forest regeneration increased,
showing a temporary regional economic depression.
4.2. Predictions of deforestation and forest
regeneration
According to our results, deforestation is positively
influenced by soil fertility and negatively influenced by
accessibility, confirming the general trends presented
by other studies in other regions and at different spatial
and temporal scales (Laurance et al., 2001; Southgate
et al., 1991; Southworth and Tucker, 2001; Veldkamp
et al., 1992; Wilson et al., 2004). Forest regeneration
shows a predominantly opposite response pattern to de-
forestation, except for the influence of forest neighbors.
The positive influence of soil fertility on deforestation
becomes weaker with time, indicating that the more
fertile soils in an area are already cleared, but also re-
flecting that further east from the Andes, the soils are
less fertile. Other factors, such as land tenure and access
to credit could further explain the encountered patterns.
The high discrimination ability of the predicted
models developed for earlier dates applied on subse-
quent periods of time, may be explained by the strong
positive influence of forest and secondary vegetation
252 A. Etter et al. / Landscape and Urban Planning 77 (2006) 240–254
neighbors on the probability of deforestation (Table 3).
Although not neglecting the contribution of many other
factors in shaping the patterns of land cover change, this
further supports the finding that, at least under the con-
ditions studied here, the spatial pattern variables, such
as proportion of forest and edge density play a major
role in the deforestation process.
4.3. Rural population
The role of population growth as a driver of de-
forestation has been frequently emphasized (Geist and
Lambin, 2001). In rural regions of Colombia with a
predominantly subsistence agricultural economy and
low use of modern agricultural technology, the expan-
sion of agricultural land is mainly driven by popula-
tion growth. This relationship is compounded by the
varying impacts of illegal markets for timber extrac-
tion, drugs and poaching. In the Caquetá region of
Colombia, the dynamics of the rural population have
been changing due to armed conflicts resulting from the
presence of illegal armies and illegal drug plantations.
In our case study, intra-regional socio-economic and
political differences can be tracked indirectly through
population movements and changing intensity of le-
gal/illegal economic activities. Population growth at
the municipality level varies according to their location
with respect to the colonization front, and the number
of immigrating or emigrating people. Our results show
a positive relation between rural population growth and
net forest cover change, reinforcing findings, such as
those of Southgate et al. (1991). In general, the munic-
ipalities located on the proper colonization front have
higher rural population growth rates with high defor-
estation rates and relatively lower rates of forest regen-
eration. There is, however, no direct relation between
the annual cleared area per person and the population
growth rate, implying that other factors, such as land
tenure, and especially, land use type, may be having an
influence. However, the lack of continuous long-term
census data common in developing countries, such as
Colombia, prevented the analysis of this trend across
the full timeframe of our study.
4.4. Conservation of biodiversity
The Caquetá region is an internationally recognized
biodiversity hotspot, making the limited knowledge
about what elements of biodiversity are being lost (due
to insufficient biological surveys) and at what rates (due
to lack of monitoring strategies) a major constraint on
conservation planning. More so, studies by Condit et
al. (2002) have shown high levels of P-diversity in the
eastern Amazon. It is urgent therefore, to inventory and
assess this diversity as part of an ecosystem mapping
program at a minimum scale of 1:100,000, thereby
providing the mapping base for evaluating what
ecosystems are being lost and where. Studies in the
Colombian Amazon and elsewhere (Duivenvoorden
and Lips, 1995; Etter, 2001; Fandiño, 1996), have
shown that ecological maps linked to a landscape
ecological approach (Zonneveld, 1995) can be used
effectively as a surrogate for biodiversity at the species
level, especially, for plants, with the advantage of
having a relatively low monetary cost.
In many cases, the knowledge required and effective
conservation measures tend to arrive too late to pro-
tect large tracts of undisturbed ecosystems adequately,
with the result that the resource base of a region gets
depleted, leaving narrower opportunities for future
conservation (Wilson et al., 2005). The rapid deforesta-
tion in biologically important areas, such as biodiver-
sity hotspots confirms this conclusion. In colonization
fronts, a trial-and-error natural resource exploitation
strategy is often the case, as most people colonizing do
not have sufficient knowledge of the area’s resources
and their ecological limitations. Governance is viewed
by Soares-Filho et al. (2004) as a force that should pro-
mote appropriate land uses to inhibit this frontier sys-
tem’s exploitative ‘entropy’, but claim that analogous
to the ‘Holling cycle’ (Gunderson and Holling, 2002),
the force of governance grows slower than the force of
exploitation. In Caquetá, the pressures of poverty, na-
tional and international markets, illegal economies and
armed conflict are effectively overruling governance.
Failure to guarantee the conservation of such biolog-
ically important areas highlights the need to look for
different alternatives. Legal conservation areas protect
such areas by acting, at least partially, as a deterrent to
land clearing (Bruner et al., 2001). This has been the
case in several areas in Colombia, such as the National
Parks of La Macarena or La Paya, which would prob-
ably be completely transformed if not protected. How-
ever, alternative approaches are needed, especially, in
areas, such as Caquetá. One possible alternative is the
initiative of the Network of Natural Reserves of the
 A. Etter et al. / Landscape and Urban Planning 77 (2006) 240–254
Civil Society (Resnatur, 2004) initiative in Colombia,
which aims at local conservation with high social in-
volvement, with close to 150 affiliated reserves cover-
ing some 50,000 ha.
Although the internal Colombian conflict has
already affected this process by slowing down the
affiliation rate, it still represents one of the best op-
portunities for conservation in highly fragmented and
dynamic landscapes, such as those detailed here. But its
success demands a socially and politically mature and
strategic-thinking environment. For areas, such as Ca-
quetá, if nothing is done, landscape transformation will
‘stabilize’ at less than 10% forest cover, with little of
the region’s rich biodiversity remaining. Conservation
scenarios in such difficult socio-economic and political
contexts should trigger national and international agen-
cies to think for more flexible and effective settings.
Acknowledgements
We acknowledge financial support to A. Etter by the
University of Queensland (GSRTA travel award) and
the Universidad Javeriana. The manuscript benefited
from useful comments provided by K. Wilson and two
anonymous reviewers.
References
Brücher, W., 1974. La Colonizacion de la Selva Pluvial en Elpiede-
monte Amazonico de Colombia. Instituto Geografico Agustin
Codazzi (IGAC), Bogota (translated from German).
Bruner, A.G., Gullison, R.E., Rice, R.E., da Fonseca, G.A.B., 2001.
Effectiveness of parks in protecting tropical biodiversity. Science
291, 125–128.
Bürgi, M., Hersperger, A.M., Schneeberger, N., 2004. Driving forces
of landscape change: current and new directions. Landsc. Ecol.
19, 857–868.
Carvajal, F., Leal, R., Molina, L.C., Cardenas, l., Diazgranados, D.,
Rodriguez, A., Etter, A., 1993. Estructura y composicion floris-
tica del bosque primario y consideraciones sobre su estado actual.
In: Malagon, D., Diazgranados, D., Saldarriaga, J.G., Rinaudo,
U. (Eds.), Aspectos Ambientales Para el Ordenamineto Territo-
rial del Occidente del Departamento del Caqueta. IGAC and the
Tropenbos Foundation, Bogota, pp. 402–531.
Clark-Labs, 2001. IDRISI 32. Idrisi Production. Clark University,
Worcester, MA.
Condit, R., Pitman, N., Leigh Jr., E.G., Chave, J., Terborgh, J., Fos-
ter, R.B., Nunez, P., Aguilar, S., Valencia, R., Villa, G., Muller-
Landau, H.C., Losos, E., Hubbell, S.P., 2002. Beta-diversity in
tropical forest trees. Science 295, 666–669.
253
Dale, V.H., O’Neill, R.V., Pedlowski, M., Southworth, R., 1993.
Causes and effects of land-use change in central Rondonia,
Brazil. Photogrammetric Eng. Remote Sens. 59, 997–1005.
Dávalos, L.M., 2001. The San Lucas mountain range in Colombia:
how much conservation is owed to the violence? Biodivers. Con-
serv. 10, 69–78.
Departamento Administrativo Nacional de Estadistica, DANE,
1993. Censo Nacional de Poblacion. http://www.dane.gov.co/
inf est/poblacion.
Duivenvoorden, J.F., Lips, J.M., 1995. A land ecological study of
soils, vegetation and plant diversity in the Colombian Amazon.
In: The Tropenbos Foundation. Wageningen, The Netherlands.
Etter, A., 1998. Mapa General de Ecosistemas de Colombia
(1:2,000,000). In: Chaves, M.E., Arango, N. (Eds.), Informe Na-
cional sobre el Estado de la Biodiversidad en Colombia, 1997.
Instituto Alexander von Humboldt, Bogota.
Etter, A. (Ed.), 2001. Punawai and Nukak: Ecological characteriza-
tion of Two National Natural Reserves in the Colombian Amazon
(Maps 1:250,000). IDEADE, Universidad Javeriana, Bogotá (in
Spanish).
Etter, A., Andrade, A., 1987. Seguimiento al proceso de coloniza-
cion en la Amazonia Colombiana. In: Proceedings of the IV
Latin American Remote Sensing Symposium, IGAC, Bogota,
pp. 84–92.
Fandiño, M.T., 1996. A Framework for Ecological Evaluation Ori-
ented at the Establishment and Management of Protected Areas:
A Case Study of the Santuario de Iguaque, Colombia. Interna-
tional Institute for Aerospace Surveys and Earth Sciences, En-
schede, The Netherlands.
Forman, R.T.T., 1995. Land Mosaics: The Ecology of Landscapes
and Regions. Cambridge, UP, NY.
Geist, H.J., Lambin, E.F., 2001. What Drives Tropical Deforestation?
A Meta-Analysis of Proximate and Underlying Causes of Defor-
estation Based on Subnational Case Study Evidence. CIACO,
Louvain-la-Neuve, http://www.geo.ucl.ac.be/LUCC/lucc.html.
Gunderson, L.H., Holling, C.S. (Eds.), 2002. Panarchy: Understand-
ing Transformations in Human and Natural Systems. Island
Press, Washington, DC.
Hernández, J., Ortiz, R., Walschburger, T., Hurtado, A., 1992. Estado
de la Biodiversidad en Colombia. CYTED, Mexico, DF.
Houghton, R.A., 1994. The worldwide extent of land-use change: in
the last few centuries, and particularly in the last several decades,
effects of land-use change have become global. Bioscience 44,
305–313.
Instituto de Estudios Ambientales, IDEAM, 2004. Estadisticas,
http://www.ideam.gov.co/publica/index4.htm. IDEAM, Minis-
terio del Medio Ambiente y Desarrollo Territorial.
Instituto Geografico Agustin Codazzi, 1983. Mapa General de Suelos
de Colombia. IGAC, Bogota.
Jensen, J.R., 1996. Introductory Digital Image Processing: A Remote
Sensing Perspective. Prentice Hall, Englewood Cliffs, NJ.
Kleinn, C., Corrales, L., Morales, D., 2002. Forest area in Costa Rica:
a comparative study of tropical forest cover estimates over time.
Environ. Monit. Assess. 73, 17–40.
Lambin, E.F., Ehrlich, D., 1997. The identification of tropical defor-
estation fronts at broad spatial scales. Int. J. Remote Sens. 18,
3551–3568.
254 A. Etter et al. / Landscape and Urban Planning 77 (2006) 240–254
Lambin, E.F., Turner II, B.L., Geist, H.J., Agbola, S.B., Angelsen,
A., Bruce, J.W., Coomes, O., Dirzo, R., Fischer, G., Folke, C.,
George, P.S., Homewood, K., Imbernon, J., Leemans, R., Li, X.,
Moran, E.E., Mortimore, M., Ramakrishnan, P.S., Richards, J.F.,
Skanes, H., Steffen, W., Stone, G., Svedin, U., Veldkamp, T.,
Vogel, C., Xu, J., 2001. The causes of landuse and land-cover
change: moving beyond the myths. Global Environ. Change 11,
261–269.
Laurance, W.F., Cochrane, M.A., Bergen, S., Fearnside, P.M., Dela-
monica, P., Barber, C., D’Angelo, S., Fernandes, T., 2001. The
future of the Brazilian Amazon. Science 291, 438–439.
Legrand, C., 1988. Colonization y Protesta Campesina en Colombia
(1850–1950). Universidad Nacional de Colombia, Bogotá.
Lucas, R., Held, A., Phinn, S., Saatchi, S., 2004. Tropical forests.
In: Ustin, S. (Ed.), Manual of Remote Sensing. Remote Sens-
ing for Natural Resource Assessment. American Society for
Photogrammetry and Remote Sensing, Bethesda, MD, 768 pp.
(Chapter 5).
Ludeke, A.K., Maggio, R.C., Reid, L.M., 1990. An analysis of an-
thropogenic deforestation using logistic regression and GIS. J.
Environ. Manage. 31, 247–259.
Malagón, D., Diazgranados, D., Saldarriaga, J.G., Rinaudo, U.
(Eds.), 1993. Aspectos Ambientales Para el Ordenamineto Ter-
ritorial del Occidente del Departamento del Caqueta. Instituto
Geografico Agustin Codazzi (IGAC) and The Tropenbos Foun-
dation, Bogota, DC.
Medlock, J., Kot, M., 2003. Spreading disease: integro-differential
equations old and new. Math. Biosci. 184, 201–222.
Mertens, B., Lambin, E.F., 1999. Modelling land cover dynamics:
integration of fine scale data with landscape attributes. Int. J.
Appl. Earth Observ. Geoinf. 1, 48–52.
Metz, C.E., 1978. Basic principles of ROC analysis. Semin. Nucl.
Med. 8, 283–298.
Myers, N., 1993. Tropical forests: the main deforestation fronts. En-
viron. Conserv. 20, 9–16.
Myers, N., Mittermeier, R.A., Mittermeier, C.G., da Fonseca, G.A.B.,
Kent, J., 2000. Biodiversity hotspots for conservation priorities.
Nature 403, 853–858.
Nagendra, H., Southworth, J., Tucker, C.J., 2003. Accessibility as a
determinant of landscape transformation in western Honduras:
linking pattern and process. Landsc. Ecol. 18, 141–158.
Nepstad, D.C., Uhl, C., Serrao, A.S., 1991. Recuperation of a de-
graded Amazonian landscape: forest recovery and agricultural
restoration. Ambio 20, 248–255.
Palacios, G. (Ed.), 2001. La Naturaleza en Disputa: Historia Ambien-
tal de Colombia, 1850–2000. Ediciones Universidad Nacional-
Colciencias, Bogota.
Phinn, S., Rowland, T., 2001. Quantifying and visualizing geometric
misregistration from landsat thematic mapper imagery band its
effect on change detection in a rapidly urbanizing catchment.
Asian Pac. Rem. Sens. J. 14, 41–54.
Pontius Jr., R.G., Batchu, K., 2003. Using the relative operating char-
acteristic to quantify certainty in prediction of location of land
cover change in India. Trans. GIS 7, 467–484.
Pontius Jr., R.G., Shusas, E., McEacherrn, M., 2004. Detecting im-
portant categorical land changes while accounting for persis-
tence. Agric. Ecosyst. Environ. 101, 251–268.
Puyravaud, J.-P., 2003. Standardizing the calculation of the annual
rate of deforestation. For. Ecol. Manage. 177, 593–596.
Resnatur, 2004. Red de Reservas naturales de la Sociedad Civil,
http://www.resnatur.org.co/.
Sanchez-Azofeifa, G., Harriss, R.C., Skole, D.L., 2001. Deforesta-
tion in Costa Rica: a quantitative analysis using remote sensing
imagery. Biotropica 33, 378–384.
Sierra, R., 2000. Dynamics and patterns of deforestation in the west-
ern Amazon: the Napo deforestation front, 1986–1996. Appl.
Geogr. 20, 1–16.
Skole, D.L., Chomentowski, W.H., Salas, W.A., Nobre, A.D., 1994.
Physical and human dimensions of deforestation in Amazonia.
Bioscience 44, 314–322.
Soares-Filho, B., Alencar, A., Nepstad, D., Cerqueira, G., Vera Diaz,
M.d.C., Rivero, S., Solorzano, L., Voll, E., 2004. Simulating the
response of land-cover changes to road paving and governance
along a major Amazon highway: the Santarem–Cuiaba corridor.
Global Change Biol. 10, 745–764.
Southgate, D., Sierra, R., Brown, L., 1991. The cause of tropical
deforestation in Ecuador: a statistical analysis. World Dev. 19,
1145–1151.
Southworth, J., Tucker, C.J., 2001. The influence of accessibil-
ity, local institutions and socioeconomic factors on forest cover
change in the mountains of western Honduras. Mt. Res. Dev. 21,
276–283.
Turner, B.L., Villar, S.C., Foster, D., Geoghegan, J., Keys, E.,
Klepeis, P., Lawrence, D., Mendoza, P.M., Manson, S., Ogneva,
H.Y., Plotkin, A.B., Salicrup, D.P., Chowdhury, R.R., Savitsky,
B., Schneider, L., Schmook, B., Vance, C., 2001. Deforestation in
the southern Yucatan peninsular region: an integrative approach.
For. Ecol. Manage. 154, 353–370.
United Nations Office for Drug Control, UNDOC, 2004. Colombia:
Coca Cultivation Survey. United Nations Office for Drug Control
and Government of Colombia, Bogota, 72 pp.
van der Hammen, M.C, 1992. El Manejo del Mundo Naturalesza y
Sociedad Entre los Yukana de la Amazonia Colombiana (Manag-
ing the World, Nature and Society by the Yukuna of the Colom-
bian Amazonia). Tropenbos-Colombia, Bogota.
Veldkamp, E., Weitz, A.M., Staritsky, I.G., Huising, E.J., 1992. De-
forestation trends in the Atlantic zone of Costa Rica: a case study.
Land Degrad. Rehabil. 3, 71–84.
Viña, A., Echavarria, F., Rundquist, D.C., 2004. Satellite change
detection analysis of deforestation rates and patterns along the
Colombia–Ecuador border. Ambio 33, 118–125.
Wiens, J., 2002. Central concepts and issues of landscape ecology.
In: Gutzwiller, K.J. (Ed.), Applying Landscape Ecology in Bio-
logical Conservation. Springer-Verlag, NY, pp. 3–21.
Wilson, K., Newton, A., Echeverria, C., Weston, C., Burgman, M.,
2004. A vulnerability analysis of the temperate forests of south
central Chile. Biol. Conserv. 122, 9–21.
Wilson, K., Pressey, R.L., Newton, A., Burgman, M., Possingham,
H., Weston, C., 2005. Measuring and incorporating vulnerability
into conservation planning. Environ. Manage., in press.
Wu, J., Hobbs, R., 2002. Key issues and research priorities in land-
scape ecology: an idiosyncratic synthesis. Landsc. Ecol. 17,
355–365.
Zonneveld, I.S., 1995. Land Ecology. Kluwer Publishers.