Biological Conservation 113 (2003) 245–256

www.elsevier.com/locate/biocon

Andean forest fragmentation and the representativeness of

protected natural areas in the eastern Andes, Colombia
D. Armenteras*, F. Gast, H. Villareal
Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, Calle 37#8-40 Mezzanine, Bogotá, Colombia

Received 1 May 2002; received in revised form 20 October 2002; accepted 8 November 2002

Abstract
Biodiversity characterization at the landscape level based on remote sensing and geographic information systems data has
become increasingly important for conservation planning. We present the results of a study of the fragmentation of Andean forests
and other ecosystems and an assessment of the representativeness at the ecosystem level of protected natural areas in the eastern
Andes of Colombia. We used satellite remote sensing data to characterize ecosystems and undertook ground truthing at six sites.
The 11 identified ecosystem types were analyzed within existing protected areas to assess the representativeness of these sites within
the region. Five ecosystems were well-represented and six of them had < 10% of their area protected. Highland ecosystems were the
best represented in protected areas due to the preponderance of highland parks in the eastern Andes. However Andean and sub
Andean forests have less than 4.5 and 6.4% of their original pre-Columbian extent currently protected. Fragmentation parameters
such as patch size, patch shape, number of patches, mean nearest neighbor distance and landscape shape index were also analyzed.
Andean, sub Andean and dry forests are highly fragmented ecosystems but there is a clear latitudinal gradient of fragmentation.
Our findings suggested that conservation efforts should be directed first toward the conservation of dry and oak forests in the center
of the eastern Andes, and then Andean and sub Andean montane forests toward the south near the border with Ecuador.
# 2003 Elsevier Science Ltd. All rights reserved.
Keywords: Colombia; Ecosystems; Andean forest; Representativeness; Fragmentation; GIS; Protected natural areas

1. Introduction tropical forests (1000–3500 m.a.s.l) are currently one of

Colombia is one of the most diverse regions for flora
and fauna in the world and has been identified as a
megadiverse country (Chaves and Arango, 1998; Fan-
diño and Ferreira, 1998). The loss of biodiversity and
landscape transformation is occurring at such a rate
that today entire ecosystem types are under threat of
disappearance (Chaves and Arango, 1998). Some esti-
mates suggest a current deforestation rate of 600,000 ha
per year (DNP, 1994). Humans have influenced the
landscape and land cover throughout the entire country.
Whilst the northernmost part of the Andes presents a
very complex geographical pattern of exceptional bio-
logical diversity (Stattersfield et al., 1998; Myers et al.,
2000) estimate that only 25% of the original tropical
forest extent remains. The northern Andean, montane
* Corresponding author. Tel.: +57-1-3406925; fax +57-1-2889564.
E-mail address: darmenteras@humboldt.org.co (D. Armenteras).
0006-3207/03/$ - see front matter # 2003 Elsevier Science Ltd. All rights reserved.
doi:10.1016/S0006-3207(02)00359-2
the major global conservation priorities due to their
biological richness, high level of endemism (Olson and
Dinerstein, 1997), and also because they are considered
amongst the least known ecosystems in the tropics
(Stadtmüller, 1987).
In Colombia, Etter (1993) suggests that only 27% of
this ecosystem’s original cover is left. With approxi-
mately 9,000,000 ha in the Andes (Etter, 1998), 40% of
which is located in the eastern slope of the eastern
Andes (IavH, 1999), the Andean montane forests are
also considered among the least known ecosystems in
the tropics (Stadtmüller, 1987). The high human popu-
lation density of the Andes adds urgency to the need for
the conservation of the last remnants of the Andean
montane forests, a conservation priority on the national
agenda (Fandiño and Ferreira, 1998).
Traditionally the biological significance of protected
areas has been evaluated by means of richness, repre-
sentativeness and vulnerability analysis (Grossman et
al., 1994). Initiatives such as the Gap analysis have been
246 D. Armenteras et al. / Biological Conservation 113 (2003) 245–256
successfully implemented in temperate zones (Scott et
al., 1989, 1991). First all the necessary information was
collected, then the degree in which biodiversity elements
are represented in a given conservation system was
evaluated (Jennings, 2000). This evaluation is usually
based on information on the percentage protected of
each type of vegetation as an estimative of its repre-
sentativeness and vulnerability (Dinerstein et al., 1995;
Stoms, 2000). Usually a figure of between 10 and 12%
of a biodiversity element present in a protected area
system is considered to be well-represented. This per-
centage, along with the number of protected areas and
their extension, are the most common indicators used to
evaluate protected systems (McNeely and Miller, 1983;
World Conservation Union, 1992; World Resources
Institute, 1994; Hummel, 1996; Noss, 1996; Duffy et al.,
1999; Pressey et al., 2002). However, in temperate zones
many initiatives of this kind are based on information
concerning the distribution of all species within an area.
In the tropics it is still difficult to use this kind of infor-
mation due to the high number of species and the lack
of knowledge of their distribution. Current tendencies
are shifting from species level evaluation towards eco-
system level (Schmidt, 1996: Hughes et al., 2000)
assuming that the higher the number of ecosystems
protected, the higher the number of species preserved
(Murray et al., 1997; Olson and Dinerstein, 1997; Stoms
et al., 1998; Noss, 1999).
Ecosystem degradation, habitat loss and fragmenta-
tion are among the principal causes of biodiversity loss
in the world (Terborgh, 1989; Whitcom et al., 1981;
Chaves and Arango, 1998; Etter, 1998). We used Geo-
graphic Information Systems and Remote Sensing tech-
nologies to make a first attempt toward analyzing the
conservation and fragmentation state of natural ecosys-
tems of the eastern Andes, an approach that has not
been undertaken previously in this part of the country.
We conducted a preliminary analysis of the representa-
tiveness of natural protected areas and ecosystem frag-
mentation analysis of the region to provide an
assessment as to the present state of the ecosystems in
this area.
We used ecosystems as an indicator of terrestrial bio-
diversity following a similar approach to Powell et al.
(2000) that used the Holdridge life zone system as their
indicator of the distribution of biodiversity in a pre-
liminary gap analysis in Costa Rica. A similar approach
has also been used in Ecuador (Sierra et al., 2002) to
assess biodiversity conservation priorities through an
analysis of ecosystem risk and representativeness.
Ecosystem fragmentation, especially in forest areas,
indicates a clear landscape change in regions with a high
human presence and has been recognized as one of the
causes of biodiversity loss (Terborgh, 1989; Whitcom et
al., 1981; Chaves and Arango, 1998). Furthermore, the
more fragmented an ecosystem is, the higher the exposure
to land use change and human pressures is. Our aim was
to provide conservationists and environmental man-
agers with information on the current state of ecosys-
tems and threats to biodiversity in the eastern Andes in
Colombia. Our analysis includes a quantitative and
descriptive analysis of the geographic distribution, the
representativeness in protected areas, and the fragmen-
tation state of natural ecosystems in the region. The
scale used for this analysis (1:250,000) is appropriate for
examining conservation priorities in the eastern Andes.
Scales between 1:250,000 and 1:500,000 are appropriate
to conduct ecosystem-level priority assessments for
small countries (or a similar extent such as the eastern
Andes) based on reliable risk and representativeness
measures taking advantage of data that is available or
easily developed (Sierra et al., 2002). Based on our
results, we propose some high-priority areas for con-
servation or establishment of special management
regimes.
2. Methods
2.1. Study area
Colombia stretches through the northwestern end of
South America between 12
260 46 N, 4
130 30 S, 66
500 54
E and 79
020 33 W. It has an area of approximately
1,142,000 km2. Colombia is a topographically variable
country. The western part is mostly mountainous as the
northern extent of the South American Andes sub-
divides into three mountain ranges when it reaches
Colombia. We centered our research in the eastern
mountain range of the Andes (the Cordillera Oriental).
This region is defined as the Colombian montane
forest ecoregion following the map of Latin America
and Caribbean ecoregions (Dinerstein et al., 1995).
However, this ecoregion excludes important zones cor-
responding to the Magdalena Valley montane forests
ecoregion, and the eastern slopes toward the Ecuador-
ian border (northern tip of the Colombian–Ecuador
northern Andean paramo and eastern Cordillera Real
montane forest ecoregions). Thus to provide a vision of
the eastern Andes as a whole, we extended the study
area to include any area above 1000 m.a.s.l. on either
side of the mountain range including the southernmost
area near the border of Ecuador. The total study area
comprised approximately 10,320,000 ha.
2.2. Ecosystems mapping
Ecosystem mapping was carried out by visual inter-
pretation of false color digital satellite imagery (12
Landsat TM scenes) corresponding to the following
years: 1989, 1991, 1992, 1994 and 1996. The procedure
delineates the different dominant ecosystem categories
 D. Armenteras et al. / Biological Conservation 113 (2003) 245–256
based on criteria such as color, texture, and context. We
cross-referenced each area with other sources of infor-
mation (e.g. refereed literature, aerial photography, field
work or other existing maps) (IGAC-ICA, 1987; IGAC,
1983; Etter, 1998; IAvH, 1999). The labeling of these
areas with categories defined by a previously defined
classification system and adding the attributes of the
interpreted individual areas to the datasets incorporated
into the geographic information system followed this.
Andean ecosystem zonation is mainly defined by alti-
tude because of its influence on temperature and oro-
graphic rainfall. A number of different classification
systems have been used in Latin America (Holdridge,
Grubb, UNESCO, IUCN) with each country adopting
its own variation on one of these systems. Generally low
elevation rainforests ( < 900–1000 m) continue to lower
montane (2300–2100 to 1200–1000 m) and upper mon-
tane (2300–2100 to 3500 m) forests. This last transition
is very important floristically because it is the upper
limit of a large number of tropical families and genera
(Van der Hammen and Hooghiemstra, 2000). The alti-
tudinal boundaries are location dependent with lowland
rainforest being separated from the montane forests by
a 900–1000 m contour line on the west flanks of the
cordillera (Doumenge et al., 1995; Gentry, 1993; Dod-
son and Gentry, 1991; Forero and Gentry, 1989; Van
der Hammen and Hooghiemstra, 2000) and by a 500 m
contour on the east flanks (Gentry, 1982, 1993). The
altitudinal limits are also clearly affected by the ‘‘Mas-
senerhebung’’ or mass-elevation effect, which causes the
occurrence of montane forest conditions at lower alti-
tudes on narrow cordilleras and outlying ridges (Flen-
ley, 1995). Sometimes a fourth altitudinal ecosystem is
found between 3000 and 3500 m: the subparamo, fol-
lowed by a grass-dominated vegetation: paramo. At the
highest altitudes permanent snow and ice caps are pre-
sent. At various elevations montane forests are subject
to frequent and/or persistent ground level cloud and are
thus termed ‘cloud forests’.
According to Hernandez (1990), a biome is defined as
an assembly of ecosystems with similar structural and
functional characteristics. The classification system
adopted in this interpretation follows the one proposed
by Hernandez and Sanchez (1987) for the higher cate-
gories (biomes). For the lower hierarchical categories
(ecosystems), the adopted classification follows that of
the General Map of Ecosystems of Colombia (Etter,
1998). To differentiate between Andean and sub Andean
montane forest, ecosystems that have important compo-
sitional and structural difference but are difficult to dis-
criminate from remote sensing data, we used GIS support
and established an artificial elevation limit of 2000 m.a.s.l.
in order to separate them. Cuatrecasas (1989) indicates
that sub Andean montane forests extend between 1000
and 2400 m.a.s.l. and the Andean forests are above the
2400 m.a.s.l. limit. Hernandez (1990) suggests these
247
limits to be around 800–1200 to 2000 m.a.s.l. for sub
Andean and above 2000–2400 m for Andean montane
forest. For the purpose of our study, we established the
lower limit at 1000 m.a.s.l. and the limit between the
two ecosystems types at around 2000 m.a.s.l.
Any kind of landscape dominated by land uses asso-
ciated with agriculture, pasture or urban sites were
assigned the category of transformed ecosystems.
We used both ERDAS Imagine (ERDAS Inc., 1999)
remote sensing processing software and the GIS soft-
ware Arcview (ESRI, 1998) to integrate all the data. As
a result of this interpretation we obtained a map of
ecosystems of the eastern mountain range at a scale of
1:250,000.
Ground testing was carried out in seven localities
(Fig. 1). Aerial photography was used to elaborate
detailed maps of around 8000 ha for each of these sites
(Fig. 2). These were used in the field to verify the satel-
lite image classification.
2.3. Representativeness
Once we had produced the ecosystem map of the
eastern Andes we overlaid it with a digitized map of the
national protected areas of this mountain range
obtained from 1:100,000 to 1:200,000 scale maps (using
only those belonging to the National Protected Areas
System). We quantified the ecosystem composition of
the protected areas and derived their representation of
the coverage as a percentage figure of the total remain-
ing in the study area. For paramos, Andean and sub
Andean forests, we also estimated the pre-transforma-
tion extent from the altitudinal range and quantified its
representativeness in terms of percentages of pre-trans-
formation extent. For other ecosystems we considered
that if 10% of the total area in each ecosystem was
protected, the ecosystem was well-represented in the
national protected areas system (McNeely and Miller,
1983; World Conservation Union, 1992; World
Resources Institute, 1994; Hummel, 1996; Noss, 1996).
This was done with the aim of obtaining a preliminary
analysis of representativeness of the protected area sys-
tem (gap analysis) using ecosystems as indicators of
terrestrial biodiversity.
2.4. Fragmentation
Based on the ecosystem map previously obtained and
with the support of GIS, an analysis of the ecosystem
fragmentation state was undertaken. The fragmentation
parameters calculated for each ecosystem type were: (1)
patch number (n) or number of fragments of a corre-
sponding ecosystem type (> 1); (2) largest patch index
(LPI) or percentage of the landscape comprised by the
largest fragment of an ecosystem type (0–100%); (3) mean
patch size (MPS) or the average size of the fragments in
248 D. Armenteras et al. / Biological Conservation 113 (2003) 245–256

Fig. 1. Ecosystem map, protected areas and ground truthed sites: [(1) 07
230 5300 N/72
230 2300 W, (2) 05
410 1800 N/73
270 4700 W, (3) 05
260 0500 N/
72
410 3000 W, (4) 05
350 1000 N / 73
250 3300 W, (5) 02
470 5100 N/74
510 1800 W and (6) 00
280 4700 N/77
170 4500 W)].

an ecosystem type ( > 0, no limit); (4) mean nearest
neighbor distance (MNND), equals the average distance
to the nearest neighboring fragment of the same eco-
system type ( > 1) and (5) landscape shape index (LSI),
the irregularity of the patch shapes (> 1, no limit).
The calculation of these indices was realized using the
software Fragstats (McGarigal and Marks, 1995). Each
one of them was chosen because of the information
provided, and the fact that they did not include redun-
dant metrics (i.e. representing the same information in
an alternate way). Each index indicates one aspect of
fragmentation, the number of patches of a particular
ecosystem might indicate that it suffers a higher rate of
disturbance (e.g. deforestation). Nevertheless, informa-
tion on the number of patches alone does not have any
interpretive value because it has no information about
area, distribution or shape of the fragments (McGarigal
and Marks, 1995), for this reason this index was calcu-
lated together with other metrics that could together be
more interpretable. Another example is the mean patch
size index, progressive reduction in the size of ecosystem
fragments is a key component of ecosystem fragmenta-
 D. Armenteras et al. / Biological Conservation 113 (2003) 245–256 249

Fig. 2. Classified map from aerial photography and ground truth transect in National Park Tamá.

tion, thus a landscape with a smaller mean patch size for
the target ecosystem than another landscape might be
considered more fragmented (McGarigal and Marks,
1995). In a similar way, the higher the mean nearest
neighbor distance the higher the fragmentation of an
ecosystem type since the distance from a patch to
another might be increasing due to human disturbances
to that ecosystem type (e.g. deforestation, land use
change, etc.). Landscape shape index reflects the shape
and complexity of the patches, higher indices indicate
higher fragmentation equal which is due to disturbances
on the edges of an ecosystem. The study area was divi-
ded into 25  25 km cells to allow for identification of
areas with a high degree of fragmentation. Each cell was
considered a landscape and the same fragmentation
indices were calculated at the landscape level in order to
compare among them. A scale was established for the
degree of fragmentation for each index scaling the ori-
ginal values into five classes (1, high to 5, low), and a
mean of all re-scaled indices was established.
250 D. Armenteras et al. / Biological Conservation 113 (2003) 245–256

3. Results the Andean xerophytic scrubs (MX) (between  1200

3.1. Ecosystem distribution
The eastern Cordillera was covered by 11 identified
natural ecosystems, which corresponded to 49%
(5,050,900 ha) of the total extension of the study area
(10,320,375 ha), transformed ecosystems accounted for
the remaining 51% of the area (5,269,475 ha). The most
extensive natural ecosystems were: the sub Andean for-
ests (17.4%), followed by the Andean montane forest
(15.2%) and the paramos (9.5%). These ecosystem
types, along with the oak forests (1.2%), correspond to
33.8% (3,492,125 ha) of the total study area (Table 1).
In terms of the forest cover, the slopes oriented towards
the Magdalena valley in the west (between 4 and 8
300
N) are more severely transformed, although there are
some extensive remnant forest patches in the mountai-
nous areas of this region.
Less degraded areas were distributed throughout the
eastern slopes of the mountain range especially between
4
N and the border with Ecuador, and also toward the
north around Cocuy National Park between 6
and 7
N.
These areas coincide with the more isolated geographic
areas where road development has been slower. However,
a clear pattern of deforestation emerged in the form of
corridors parallel to communication roads and main riv-
ers. Nevertheless, forests were still distributed in a con-
tinuous elevation gradient south of the eastern Andes.
Ecosystems with a restricted geographic distribution
(mainly due to local climate and soil conditions) such as
and 2600 m.a.s.l.) covered 1.25% (128,700 ha) of the
study area. They were in specific areas of Abrego-Ocaña
(north of Santander); Chicamocha canyon (Boyacá);
Soacha and Guatavita (Cundinamarca); Villa de Leyva
(Boyacá); valleys of the rivers Guaitara, Juanambú
(Nariño), Guachicono (Cauca), and Cabrera (Huila).
Dry forest and secondary xerophytic scrubs (BS) were
found below 1200 m.a.s.l. and occupied 3.7% of the
area (377,475 ha). This ecosystem type was also geo-
graphically restricted to areas of the Chicamocha can-
yon (Santander), around Cúcuta (north of Santander)
and in the Patı´a (Cáuca-Nariño), Cabrera (Huila) and
Negro (Cundinamarca) river valleys and to the north in
the mountainous area of Perijá (Cesar-Guajira).
All six sites visited for ground truthing had Andean
forests and paramos as their main ecosystems. The field
visit confirmed that the image classification was correct
in those sites although the degree of forest fragmenta-
tion that was appreciated in one of the sites from the
photographs was, in fact, higher than the impression
obtained from satellite imagery, this was mainly due to
the scale of work. The southern windows showed clearly
that the natural cover shows no sign of degradation.
3.2. Representativeness
The protected natural ecosystems of the eastern Andes
cover an area of 830,555 ha (8.05% of study area); 44,225
ha within this area have already been transformed (over
5% of the protected area) and patterns of land use

Table 1
Current coverage of natural ecosystems in the eastern Andes and percentage of protection in the system of protected areas

Natural ecosystem type Original Current Study Original Protected Original Total Current Current
area area area area (ha) area protected ecosystem ecosystem
(ha) (ha) (%) remaining protected area (%) area area
(%) (%) protected unprotected
(%) (%)

Dry forest and secondary naa 377,475 3.7 na 851 na 0.1 0.2 99.8
xerophytic shrubs ( < 1200 m)*
Xerofic shrubs of andean na 128,700 1.2 na 0 na 0 0 100
localities ( > 1200 m)*
Sub-andean montane forests 3,978,925 1,796,250 17.4 45% 254,125 6.4% 30.6 14.1 85.9
(1,000 – 2,000 m )
Andean montane forests 3,812,775 1,567,525 15.2 41% 173,550 4.5% 20.9 11.2 88.8
(2,000 – 3000–3500 m)
Humid paramos na 920,875 8.9 na 322,075 na 38.8 35 65
Dry paramos na 60,275 0.6 na 850 na 0.1 1.4 98.6
Superparamo na 20,925 0.2 na 20,925 na 2.5 100 0
Snow na 3775 0.0 na 3775 na 0.5 100 0
Oak forests na 128,350 1.2 na 10,175 na 1.2 7.9 92.1
Intra Andean savanahs na 29,950 0.3 na 0 na 0 0 100
Wetlands na 16,800 0.2 na 0 na 0 0 100

Total 5,050,900 48.9 786,326 94.7

*Ecosystems with a high level of degradation.

a
na, Not available.
 D. Armenteras et al. / Biological Conservation 113 (2003) 245–256 251

change within the borders of some protected areas can be Table 2

appreciated. The remaining 786,326 ha are natural eco- Ecosystem composition of the 11 protected areas belonging to the
national parks system of the eastern Andes, Colombia
systems, equivalent to 15.6% of the existing natural eco-
systems in the mountain range (4,862,925 ha). We only Protected area Ecosystem Area
considered the areas above 1000 m in our calculations.
(ha) %
In general terms, the representation objective of 10%
is not met for six ecosystems. Dry and humid paramos LOS ESTORAQUES Dry forest 851 100.0
(highland ecosystems) had a high percentage of their PNN CHINGAZA Humid Paramos 34,375 71.5
Subandean forest 2200 4.6
area protected with a total extent of 322,925 ha, which
Andean forest 11,100 23.1
equates to 36.4% of its total area within the study area Transformed 375 0.8
(Table 1). Paramos were followed by Andean montane PNN CORDILLER Humid Paramos 3800 2.7
forest (both Andean and sub Andean) in the degree of LOS PICACHOS
protection with 427,675 ha protected (25.3% of the Subandean forest 119,150 84.7
Andean forest 16,200 11.5
existing total). However, only 41 and 45% of these two
Transformed 1575 1.1
Andean ecosystems remain from their original extent PNN CUEVA DE Subandean forest 6050 82.3
(see Table 1) and the percentage of original pre-trans- LOS GUACHAROS
formed protected area is 4.5% for Andean and 6.4% for Andean forest 1300 17.7
sub Andean forests. PNN EL COCUY Humid Paramos 122,375 40.8
Subandean forest 73,775 24.6
In contrast, oak forests had only 7.9% of their current
Andean forest 68,625 22.9
surface protected (10,175 ha) in the interior of Sanctu- Superparamo 20,925 7.0
aries of Flora and Fauna de Iguaque and the Guanentá- Nival 3775 1.3
Alto Fonce river, with a total extension of 128,350 ha. Transformed 10,675 3.6
Dry ecosystems were much less protected than oak for- PNN PISBA Dry Paramos 20,800 58.6
Andean forest 5200 14.7
ests. Only about 0.1% of these ecosystems were pro-
Transformed 9475 26.7
tected in the Estoraques area. Over 90% of the PNN SUMAPAZ Dry Paramos 129,375 59.3
protected ecosystems corresponded to highland ecosys- Subandean forest 22,000 10.1
tems and over one third of them were paramos. Within Andean forest 62,275 28.6
each protected area, 9 out of 11 protected zones con- Transformed 4350 2.0
PNN TAMA Dry Paramos 5675 10.4
tained this ecosystem type and in five of them it con-
Subandean forest 27,125 49.6
stituted more than two thirds of their area (Table 2). Andean forest 8850 16.2
Some parks had an elevation gradient, which was Transformed 13,050 23.9
reflected in the diversity and extension of the ecosys- SFF GALERAS Dry Paramos 3125 38.6
tems, and their limits extend beyond the defined 1000 Andean forest 3825 47.2
Transformed 1150 14.2
m.a.s.l. lower study area limit (Cocuy, Sumapaz and
PNN GUANENTA Oak Forests 7325 74.2
Tamá National Parks). The remaining parks were char- Humid Paramos 2550 25.8
acterized by the dominance of a single ecosystem. SFF IGUAQUE Oak Forests 2850 39.2
Unfortunately, and despite their legal protection status, Dry Paramos 850 11.7
some parks showed remarkable levels of human inter- Transformed 3575 49.1
vention that were clear from the percentage of ecosys-
tems transformed found within their limits. This reflects
the fact that a declaration of a protected area does not Table 3
Fragmentation indices of the eastern Andes ecosystems, Colombiaa
guarantee its protection: 7 out of 11 parks analyzed
show some degree of land use change due to human Ecosystem LPI NP MPS LSI MNND
activities.
Dry forests 1.075 135 2796.1 11.749 1446.4
Xerofic shrubs 0.288 16 8043.7 10.82 15,444.4
3.3. Fragmentation Subandean forests 2.929 302 5947.8 18.175 1301.3
Andean forests 5.081 118 13,177.9 19.162 1602.7
The most fragmented ecosystems corresponded to the Humid Paramos 4.514 40 23,021.8 14.998 6374.1
Dry Paramos 0.398 6 10,045.8 10.618 7859.6
Andean montane forests, the sub Andean montane for-
Superparamo 0.157 3 6975 10.383 4626.9
ests and the dry forests (Table 3). The Andean forests Nival 0.026 4 943.7 10.231 926.7
had 118 fragments, however they had the largest patch Oak forests 0.426 19 6755.2 11.37 1566.8
index of 5, meaning that a single fragment of this eco- Intra andean savanahs 0.113 4 7487.5 10.31 3177
system occupied 5% of the total of the studied area. In Wetlands 0.054 7 2400 10.318 98,307.9
the sub Andean montane forest the number of frag- a
LPI, largest patch index; NP, number of patches; MPS, mean
ments was 302 and the largest patch index was 3%. The patch size; LSI, landscape shape index; MNND, mean nearest neigh-
mean nearest neighbor distance was similar in the bour distance.
252 D. Armenteras et al. / Biological Conservation 113 (2003) 245–256
Andean and the sub Andean montane forests: 1.3 and
1.6 km. There were some differences in the mean patch
size: 5947 and 13,177 ha. Dry forests also appeared
fragmented, with 135 fragments, a mean patch size of
2796 ha, a mean nearest neighbor distance of 1.4 km
and a largest patch index of 1.08%. Nevertheless, this
fragmentation changed over the latitudinal gradient.
There were some areas dominated by a matrix of trans-
formed landscapes in which dispersed fragments of the
original ecosystems occasionally appeared. The area
between 4
and 8
300 N concentrated the greater num-
ber of patches of Andean and sub Andean forests. A
smaller number of fragments, but with a continuous and
extended presence of forest were found around the
Cocuy National Park and the in the south, toward the
border with Ecuador. This is why the largest patch
index values appeared to be relatively high in these
ecosystems.
From the results of the 25  25 km analysis, the 30
cells with lower level of fragmentation were geo-
graphically located around five protected areas: Cocuy,
Pisba, Sumapaz, Tama and Picachos (Fig. 3). The rest
were in the southern Andes, where there is only one
protected area established, the Cueva de los Guacharos.
This part of the eastern Andes is a very interesting area
for conservation because ecosystems are less altered,
there is a clear continuous elevation gradient of forest
cover toward the Amazonia (i.e. influence of Amazo-
nian ecosystems and species) and high biological rich-
ness (IavH, 1999).
4. Discussion
Transformed ecosystems covered 51% of the total
study area. The other 49% corresponded to natural
ecosystems, such as paramos, and Andean and sub
Andean forests. These ecosystems were the best repre-
sented in protected areas of the eastern Cordillera (35,
14.1 and 11%, respectively). This is due to the pre-
ponderance of highland parks in the Andes which ori-
ginated in the establishment of Colombian protected
natural areas without organized planning. However,
when incorporating criteria based only on the percen-
tage of total current area of an ecosystem, we are failing
to consider the important reduction dating from pre-
Columbian times. If we do take this into account, there
is only a 41% and 45% of the original pre-transformed
extent of Andean and sub Andean forest left and the
percentage of protected area drop to figs. of 4.5% for
Andean and 6.4% for sub Andean forests. Further,
despite the fact that we considered 10% of the total
current area of an ecosystem as the limit to define whe-
ther or not an ecosystem is well protected within the
national areas protected system, this is more a political
conservation target than a scientifically based result
(Soulé and Sanjayan, 1998). Furthermore the declara-
tion of a protected area does not guarantee its protec-
tion, 7 of 11 national parks analyzed show some degree
of transformation due to human activities and inter-
active management practices might be needed.
Despite the high level of degradation, threat of habi-
tat loss and further degradation and recognized scien-
tific interest, neither dry nor oak forests were properly
represented in the current protected areas system of the
region. We recommend that the conservation of the
remaining fragments and the recovery of secondary
vegetation in these areas be made a priority. Due to the
scale and approach we used in this research, only two
areas of this ecosystem were identified as high priority
for conservation: the Soacha-Bojacá region and the area
around the protected area Estoraques (Fig. 4). Further
work on a more detailed scale will allow better assess-
ment of the real distribution of dry ecosystem remnants
and the degree of anthropogenic disturbance. We sug-
gest further protection of oak forest areas around the
Sanctuaries of Flora and Fauna de Iguaque, and the
Guanentá-Alto Fonce river should also be considered
high priority areas for conservation (Fig. 4).
The area between 2
N and the border with Ecuador
(Bota Caucana and Nudo de los Pastos) was also iden-
tified as one with the greatest interest for conservation
(Fig. 4). These areas are in a better condition, have
lower degree of fragmentation and have wide elevation
gradients of continuous forest cover and are not yet
protected. Consequently, conservation actions should
be directed toward this sector. Other identified areas of
interest are located in areas already within the national
park system (Cocuy, Tamá, Sumapaz and Picachos
mountain range). The central region of the eastern
Andes has the highest degree of ecosystem alteration
and the fewest remaining fragments.
Some of the fragmentation values obtained in the
remaining ecosystems owe more to their natural geo-
graphic distribution and topographic landscape hetero-
geneity than to disturbances caused by human action.
This is the case for dry forests that were, to a certain
degree, naturally fragmented at this regional level
because they were located in specific soil and climatic
conditions. Paramos and wetlands show similar restric-
ted distributions. The mean neighbor distance and mean
patch size of paramos was high because of its natural
location in the highest mountains. Dry forest frag-
mented areas correspond to the Chicamocha canyon.
Dry ecosystems have a high level of degradation and
fragmentation that are not easy to differentiate at this
scale of research when observed from satellite images.
These are only preliminary results that suggest
conservation actions in specific ecosystems and areas.
However, it is necessary to consider the inclusion of
other types of land management for conservation
practices outside the national parks system, such as
 D. Armenteras et al. / Biological Conservation 113 (2003) 245–256 253

Fig. 3. The degree of fragmentation, 25  25 km division of the eastern Andes, Colombia and protected areas in the zone (PNN, Natural National
Park).

indigenous territories, private reserves and forest
reserves. These areas were not incorporated because
of the scale of work we used in this study. The
incorporation of these areas in later analyses at a
more detailed scale (1:100,000 or 1:25,000) will help
obtain more specific information to identify a port-
folio of sites for conservation at different geographic
levels.
We also suggest redefining some of the protected
areas of the national parks system to include surround-
ing areas of natural ecosystems before they are further
degraded, and in the mid term conduct an analysis of the
254 D. Armenteras et al. / Biological Conservation 113 (2003) 245–256

Fig. 4. Ecosystems and suggested areas for conservation action of the eastern Andes, Colombia.

effective protection level of each area. There is also a
need to clearly quantify land use change and deforesta-
tion rates through a multitemporal evaluation based on
remote sensing images. This multitemporal evaluation of
change will become a valuable criterion for defining the
degree of threat to ecosystems coupled with other factors
of threat such as infrastructure and population growth.
We consider this analysis an intermediate step between
using a coarse approach (Dinerstein et al., 1995) and a
detailed species gap analysis (Scott et al., 1991). We also
expect to be able to incorporate species distribution maps
to determine areas of high species richness and endemism
when more taxonomic data becomes available.
Acknowledgements
Thanks to C. Franco for her contribution to this work
and to M. Mulligan, S. Newey and the reviewers for
 D. Armenteras et al. / Biological Conservation 113 (2003) 245–256
their comments on the manuscript. This work was par-
tially supported by The Nature Conservancy and Fun-
dación Natura, Colombia.
References
Chaves, M.E., & Arango, N. (Eds.) (1998). Informe nacional sobre el
estado de la biodiversidad 1997. Instituto de Investigación de
Recursos Biológicos Alexander von Humboldt, PNUMA and Min-
isterio de Medio Ambiente. 3 vol. Bogotá, Colombia.
Cuatrecases, J., 1989. Aspectos de la vegetación natural en Colombia.
Rev. Pérez Arbelaezia. 11, 155–287. Bogotá, Colombia..
Departamento Nacional de Planeación, 1994. Polı́tica ambiental.
CONPES. Presidencia de la República de Colombia, Bogota,
Colombia.
Dinerstein, E., Olson, D.M., Graham, D.H., Webster, A.L., Primm,
S.A., Bookbinder, M.P., Ledec, G., 1995. A Conservation Assess-
ment of the Terrestrial Ecoregions of Latin America and the Car-
ibbean. World Wildlife Fund and World Bank, Washington DC,
USA.
Dodson, C.H., Gentry, A.H., 1991. Biological extinction in western
Ecuador. Ann. Missouri Bot. Garden 78, 273–295.
Doumenge, C., Gilmour, D, Ruiz Pérez, M., Blockhus, J., 1995. Tropi-
cal montane cloud forests: conervation status and management issues.
In: Hamilton, L.S., Juvik, J.O., Scatena, F.N. (Eds.), Tropical
montane cloud forest. Springer-Verlag, New York, USA, pp. 24–37.
Duffy, D.C., Boggs, K., Hagenstein, R.H., Lipkin, R.y., Michaelson,
J.A., 1999. Landscape assessment of the degree of protection of
Alaska’s terrestrial biodiversity. Conservation Biology 13 (6), 1332–
1343.
Erdas Inc, 1999. Erdas Imagine Version 8.4. ERDAS, Inc, Atlanta,
Georgia, USA.
Esri Inc, 1998. Arview GIS Version 3.1. Environmental Systems
Research Institute, USA.
Etter, A., 1993. Diversidad ecosistemica en Colombia hoy. In: Cárde-
nas, S., Correa, H.D. (Eds.), Nuestra Diversidad Biológica. Funda-
ción Alejandro Escobar, Colección Marı́a Restrepo de Angel.
CEREC, Bogotá, Colombia.
Etter, A., 1998. Mapa general de ecosistemas de Colombia
(1:1.500.000). In: Chaves, M.E., Arango, N. (Eds.), Informe nacio-
nal sobre el estado de la biodiversidad 1997. Instituto de Investiga-
ción de Recursos Biológicos Alexander von Humboldt, PNUMA
and Ministerio de Medio Ambiente. 3 vol, Bogotá, Colombia.
Fandiño, M.C., Ferreira, P. (Eds.), 1998. Colombia biodiversidad
siglo XXI: propuesta técnica para la formulación de un plan de
acción nacional en biodiversidad. Instituto de Investigación de
Recursos Biológicos Alexander von Humboldt, Ministerio del
Medio Ambiente y Departamento Nacional de Planeación, Bogotá,
Colombia.
Flenley, J.R., 1995. Cloud forest, the Massenerhebung effect, and
ultraviolet insolation. In: Hamilton, L.S., Juvik, J.O., Scatena, F.N.
(Eds.), Tropical Montane Cloud Forests. Ecol. Studies (Vol. 110).
Springer-Verlag, New York, pp. 150–155.
Forero, E., Gentry, A.H., 1989. Lista anotada de las plantas del
Departamento del Chocó, Colombia. Biblioteca J.J. Triana No. 10.
Instituto de Ciencias Naturales, Museo de Historia Natural, Uni-
versidad Nacional de Colombia, Bogotá.
Gentry, A.H., 1982. Neotropical floristic diversity: phytogeographical
connections between Central and South America, Pleistocene cli-
matic fluctuations, or an accident of the Andean orogeny? Ann.
Missouri Bot. Garden 69, 557–593.
Gentry, A.H. 1993. Overview of the Perúvian flora. In Brako, L.,
Zarucchi, J.L., Catalogue of the flowering plants and gymnosperms
of Perú. Monogr. Syst. Bot. Missouri Bot. Garden 45: xxix–xl.
255
Grossman, D. H., Goodin, K.L. & Reuss, C.L. (Eds.) (1994). Rare
plant communities of the coterminous United States—an initial
survey. Prepared for the USDI Fish and Wildlife Service. The Nat-
ure Conservancy, Arlington, VA.
Hernández, J., 1990. La selva en Colombia. In: Carrizoza, J., Her-
nández, J. (Eds.), Selva y Futuro. El Sello Editorial, Bogotá,
Colombia, pp. 28–50.
Hernández, J., Sanchez, E., 1987. Ensayo preliminar sobre los biomas
terrestres de Colombia. In: Sánchez, E., Hernández, J., Rueda, V.
(Eds.), Nuevos parques naturales. Instituto de Recursos Naturales
Renovables, Bogotá, Colombia.
Hughes, J., Daily, G., Ehrlich, P., 2000. Conservation of insect
diversity: a habitat approach. Conservation Biology 14 (6), 1788–
1797.
Hummel, M., 1996. Protecting Canada’s endangered species: an own-
er’s manual. Key Porter, Toronto, Canada.
IavH, 1999. Caracterización de la biodiversidad en áreas prioritarias
de la vertiente oriental de la cordillera Oriental. Instituto de Inves-
tigación de Recursos Biológicos Alexander von Humboldt, Villa de
Leyva, Colombia.
IGAC, 1983. Bosques de Colombia (escala 1:500.000). Instituto Geo-
gráfico Agustı́n Codazzi, Bogotá, Colombia.
IGAC-ICA, 1987. Mapa de Uso actual del suelo en Colombia (escala
1:500.000). Instituto Geográfico Agustı́n Codazzi, Instituto Colom-
biano Agropecuario, Bogotá, Colombia.
Jennings, M.D., 2000. Gap analysis: concepts, methods, and recent
results. Landscape Ecology 15, 5–20.
Mcgarigal, K., Marks, B.J., 1995. Fragstats: spatial pattern analysis
program for quantifying landscape structure. Gen. Tech. Rep.
PNW-GTR-351. US Departament of Agriculture, Forest Service,
Pacific Northwest Research Station, USA, Portland, OR.
McNeely, J.A., Miller, K.R., 1983. National parks and protected
areas. UN Economic and Social Commission for Asia and the
Pacific, Bangkok.
Murray, M., Green, M., Bunting, G, Paines, J., 1997. Priorities for
biodiversity conservation in the tropics. WCMC Biodiversity Series
6. World Conservation Press, Cambridge UK.
Myers, N., Mittermeier, R.A., Mittermeier, C.G., Da Fonseca,
G.A.B., Kent, J., 2000. Biodiversity hotspots for conservation prio-
rities. Nature 403, 853–858.
Noss, R.F., 1996. In: Wright, R.G. (Ed.), National parks and pro-
tected areas: their role in environmental protection. Blackwell
Science, Cambridge, USA, pp. 91–119.
Noss, R.F., 1999. Assessing and monitoring forest biodiversity: a
suggested framework and indicators. Forest Ecology and Manage-
ment 111, 135–146.
Olson, D.M., Dinerstein, E., 1997. Global 2000: conserving the
world’s distinctive ecoregions. WWF-US, USA.
Powell, G.V.N., Barborak, J., Rodriguez, M., 2000. Assessing repre-
sentativenes of protected natural areas in Costa Rica for conserving
biodiversity: a preliminary gap analyis. Biological Conservation 93,
35–41.
Pressey, R.L., Whish, G., 2002. L, Barrett, T. W., Watts, M.E. Effec-
tivenes of protected areas in north-eastern New South Wales: recent
trends in six measures. Biological Conservation 106, 57–69.
Schmidt, K., 1996. Rare habitat vie for protection. Science 274, 916–
918.
Scott, J.M., Csuti, B., Estes, J.E., Anderson, H., 1989. State assess-
ment of biodiversity protection. Conservation Biology 3, 85–87.
Scott, J.M., Csuti, B., Estes, J.E., Caicco, S., 1991. Gap analysis of
species richness and vegetation cover: an integrated biodiverstiy
conservation strategy. In: Kohm, K. (Ed.), Balancing on the brink
of extinction: the endangered species act and lessons for the future.
Island Press, Washington DC USA, pp. 282–297.
Sierra, R., Campos, F., Chamberlin, J., 2002. Assessing biodiversity
conservation priorities: ecosystem risk and representativeness in
continental Ecuador. Landscape and Urban Planning 59, 95–110.
256 D. Armenteras et al. / Biological Conservation 113 (2003) 245–256
Soulé, M.E., Sanjayan, M.A., 1998. Conservation targets: do they
help? Science 279, 2060–2061.
Stadtmüller, T., 1987. Cloud forests in the humid tropics: a biblio-
graphic review. Centro Agronómico Tropical de Investigación y
Enseñanza, Turrialba, Costa Rica.
Stattersfield, A.J., Crosby, M.J, Long, A.J., Wege, D.C., 1998. Ende-
mic bird areas of the world: priorities for biodiversity conservation.
BirdLife Conservation Serires, Cambridge.
Stoms, D., 2000. Gap management status and regional indicators of
threats to biodiversity. Landscape Ecology 15, 5–20.
Stoms, D.M., Borchert, M.I., Church, R.L., 1998. A systematic process
for selecting representative research natural areas. Natural Areas
Journal 18 (4), 338.
Terborgh, J., 1989. Where have all the birds gone? Princeton Uni-
versity Press, USA, New Jersey.
Van der Hammen, T., Hooghiemstra, H., 2000. Neogene and
Quaternary history of vegetation, climate, and plant diversity in
Amazonia. Quaternary Science Reviews 19, 725–742.
Whitcom, R.F., Robbins, C.S., Lynch, J.F., 1981. Effects of forest
fragmentation on avifauna of the estern deciduous forest. In: Burgess,
R.L., Sharpe, D.M. (Eds.), Forest island dynamics in a man-domi-
nated landscapes. Springer-Verlag, New York, USA, pp. 125–205.
World Conservation Union. 1992. IUCN Bulletin 43.
World Resources Institute, 1994. World resources, 1994–1995. Oxford
University Press, New York, USA.