Biological Conservation 113 (2003) 245256 www.elsevier.

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Andean forest fragmentation and the representativeness of protected natural areas in the eastern Andes, Colombia
D. Armenteras*, F. Gast, H. Villareal
n de Recursos Biolo gicos Alexander von Humboldt, Calle 37#8-40 Mezzanine, Bogota , Colombia Instituto de Investigacio 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 identied 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 ndings suggested that conservation eorts should be directed rst 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 Colombia is one of the most diverse regions for ora and fauna in the world and has been identied as a megadiverse country (Chaves and Arango, 1998; Fandin o 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 estimates suggest a current deforestation rate of 600,000 ha per year (DNP, 1994). Humans have inuenced the landscape and land cover throughout the entire country. Whilst the northernmost part of the Andes presents a very complex geographical pattern of exceptional biological diversity (Statterseld 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).

tropical forests (10003500 m.a.s.l) are currently one of 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 (Stadtmu ller, 1987). In Colombia, Etter (1993) suggests that only 27% of this ecosystems original cover is left. With approximately 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 (Stadtmu ller, 1987). The high human population 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 (Fandin o and Ferreira, 1998). Traditionally the biological signicance of protected areas has been evaluated by means of richness, representativeness and vulnerability analysis (Grossman et al., 1994). Initiatives such as the Gap analysis have been

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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 representativeness and vulnerability (Dinerstein et al., 1995; Stoms, 2000). Usually a gure of between 10 and 12% of a biodiversity element present in a protected area system is considered to be well-represented. This percentage, 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; Duy 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 dicult to use this kind of information due to the high number of species and the lack of knowledge of their distribution. Current tendencies are shifting from species level evaluation towards ecosystem 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 fragmentation 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 Geographic Information Systems and Remote Sensing technologies to make a rst attempt toward analyzing the conservation and fragmentation state of natural ecosystems 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 representativeness of natural protected areas and ecosystem fragmentation 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 biodiversity 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 preliminary 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 managers with information on the current state of ecosystems 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 fragmentation 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 conservation 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 subdivides 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 dened 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 corresponding to the Magdalena Valley montane forests ecoregion, and the eastern slopes toward the Ecuadorian border (northern tip of the ColombianEcuador 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 interpretation 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 dierent dominant ecosystem categories

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based on criteria such as color, texture, and context. We cross-referenced each area with other sources of information (e.g. refereed literature, aerial photography, eld work or other existing maps) (IGAC-ICA, 1987; IGAC, 1983; Etter, 1998; IAvH, 1999). The labeling of these areas with categories dened by a previously dened classication 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 dened by altitude because of its inuence on temperature and orographic rainfall. A number of dierent classication 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 ( < 9001000 m) continue to lower montane (23002100 to 12001000 m) and upper montane (23002100 to 3500 m) forests. This last transition is very important oristically because it is the upper limit of a large number of tropical families and genera (Van der Hammen and Hooghiemstra, 2000). The altitudinal boundaries are location dependent with lowland rainforest being separated from the montane forests by a 9001000 m contour line on the west anks of the cordillera (Doumenge et al., 1995; Gentry, 1993; Dodson and Gentry, 1991; Forero and Gentry, 1989; Van der Hammen and Hooghiemstra, 2000) and by a 500 m contour on the east anks (Gentry, 1982, 1993). The altitudinal limits are also clearly aected by the Massenerhebung or mass-elevation eect, which causes the occurrence of montane forest conditions at lower altitudes on narrow cordilleras and outlying ridges (Flenley, 1995). Sometimes a fourth altitudinal ecosystem is found between 3000 and 3500 m: the subparamo, followed by a grass-dominated vegetation: paramo. At the highest altitudes permanent snow and ice caps are present. 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 dened as an assembly of ecosystems with similar structural and functional characteristics. The classication system adopted in this interpretation follows the one proposed by Hernandez and Sanchez (1987) for the higher categories (biomes). For the lower hierarchical categories (ecosystems), the adopted classication follows that of the General Map of Ecosystems of Colombia (Etter, 1998). To dierentiate between Andean and sub Andean montane forest, ecosystems that have important compositional and structural dierence but are dicult to discriminate from remote sensing data, we used GIS support and established an articial 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

limits to be around 8001200 to 2000 m.a.s.l. for sub Andean and above 20002400 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 associated 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 software 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 eld to verify the satellite image classication. 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 quantied the ecosystem composition of the protected areas and derived their representation of the coverage as a percentage gure of the total remaining in the study area. For paramos, Andean and sub Andean forests, we also estimated the pre-transformation extent from the altitudinal range and quantied its representativeness in terms of percentages of pre-transformation 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 system (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 corresponding ecosystem type ( > 1); (2) largest patch index (LPI) or percentage of the landscape comprised by the largest fragment of an ecosystem type (0100%); (3) mean patch size (MPS) or the average size of the fragments in

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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 ecosystem 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 redundant 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 suers a higher rate of disturbance (e.g. deforestation). Nevertheless, information 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 calculated 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-

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. Fig. 2. Classied map from aerial photography and ground truth transect in National Park Tama

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 reects 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 divided into 25 25 km cells to allow for identication 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 original values into ve classes (1, high to 5, low), and a mean of all re-scaled indices was established.

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3. Results 3.1. Ecosystem distribution The eastern Cordillera was covered by 11 identied 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 forests (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 mountainous 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 rivers. Nevertheless, forests were still distributed in a continuous elevation gradient south of the eastern Andes. Ecosystems with a restricted geographic distribution (mainly due to local climate and soil conditions) such as

the Andean xerophytic scrubs (MX) (between $ 1200 and 2600 m.a.s.l.) covered 1.25% (128,700 ha) of the study area. They were in specic areas of Abrego-Ocana ); (north of Santander); Chicamocha canyon (Boyaca Soacha and Guatavita (Cundinamarca); Villa de Leyva ); valleys of the rivers Guaitara, Juanambu (Boyaca (Narino), 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 geographically restricted to areas of the Chicamocha cancuta (north of Santander) yon (Santander), around Cu uca-Narino), Cabrera (Huila) and and in the Pata (Ca Negro (Cundinamarca) river valleys and to the north in (Cesar-Guajira). the mountainous area of Perija All six sites visited for ground truthing had Andean forests and paramos as their main ecosystems. The eld visit conrmed that the image classication was correct in those sites although the degree of forest fragmentation 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 area (ha) Current area (ha) Study area (%) Original area remaining (%) na na 45% 41% na na na na na na na Protected (ha) Original area protected (%) na na 6.4% 4.5% na na na na na na na Total protected area (%) Current ecosystem area protected (%) 0.2 0 14.1 11.2 35 1.4 100 100 7.9 0 0 Current ecosystem area unprotected (%) 99.8 100 85.9 88.8 65 98.6 0 0 92.1 100 100

Dry forest and secondary xerophytic shrubs ( < $ 1200 m)* Xeroc shrubs of andean localities ( > $ 1200 m)* Sub-andean montane forests (1,000 2,000 m ) Andean montane forests (2,000 $ 30003500 m) Humid paramos Dry paramos Superparamo Snow Oak forests Intra Andean savanahs Wetlands Total

naa na 3,978,925 3,812,775 na na na na na na na

377,475 128,700 1,796,250 1,567,525 920,875 60,275 20,925 3775 128,350 29,950 16,800 5,050,900

3.7 1.2 17.4 15.2 8.9 0.6 0.2 0.0 1.2 0.3 0.2 48.9

851 0 254,125 173,550 322,075 850 20,925 3775 10,175 0 0 786,326

0.1 0 30.6 20.9 38.8 0.1 2.5 0.5 1.2 0 0 94.7

*Ecosystems with a high level of degradation. a na, Not available.

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change within the borders of some protected areas can be appreciated. The remaining 786,326 ha are natural ecosystems, equivalent to 15.6% of the existing natural ecosystems in the mountain range (4,862,925 ha). We only considered the areas above 1000 m in our calculations. In general terms, the representation objective of 10% is not met for six ecosystems. Dry and humid paramos (highland ecosystems) had a high percentage of their area protected with a total extent of 322,925 ha, which equates to 36.4% of its total area within the study area (Table 1). Paramos were followed by Andean montane forest (both Andean and sub Andean) in the degree of protection with 427,675 ha protected (25.3% of the existing total). However, only 41 and 45% of these two Andean ecosystems remain from their original extent (see Table 1) and the percentage of original pre-transformed protected area is 4.5% for Andean and 6.4% for sub Andean forests. In contrast, oak forests had only 7.9% of their current surface protected (10,175 ha) in the interior of Sanctuaries of Flora and Fauna de Iguaque and the Guanenta Alto Fonce river, with a total extension of 128,350 ha. Dry ecosystems were much less protected than oak forests. Only about 0.1% of these ecosystems were protected in the Estoraques area. Over 90% of the protected ecosystems corresponded to highland ecosystems and over one third of them were paramos. Within each protected area, 9 out of 11 protected zones contained this ecosystem type and in ve of them it constituted more than two thirds of their area (Table 2). Some parks had an elevation gradient, which was reected in the diversity and extension of the ecosystems, and their limits extend beyond the dened 1000 m.a.s.l. lower study area limit (Cocuy, Sumapaz and National Parks). The remaining parks were charTama acterized by the dominance of a single ecosystem. Unfortunately, and despite their legal protection status, some parks showed remarkable levels of human intervention that were clear from the percentage of ecosystems transformed found within their limits. This reects the fact that a declaration of a protected area does not guarantee its protection: 7 out of 11 parks analyzed show some degree of land use change due to human activities. 3.3. Fragmentation The most fragmented ecosystems corresponded to the Andean montane forests, the sub Andean montane forests and the dry forests (Table 3). The Andean forests had 118 fragments, however they had the largest patch index of 5, meaning that a single fragment of this ecosystem occupied 5% of the total of the studied area. In the sub Andean montane forest the number of fragments was 302 and the largest patch index was 3%. The mean nearest neighbor distance was similar in the

Table 2 Ecosystem composition of the 11 protected areas belonging to the national parks system of the eastern Andes, Colombia Protected area Ecosystem Area (ha) LOS ESTORAQUES PNN CHINGAZA Dry forest Humid Paramos Subandean forest Andean forest Transformed Humid Paramos Subandean forest Andean forest Transformed Subandean forest Andean forest Humid Paramos Subandean forest Andean forest Superparamo Nival Transformed Dry Paramos Andean forest Transformed Dry Paramos Subandean forest Andean forest Transformed Dry Paramos Subandean forest Andean forest Transformed Dry Paramos Andean forest Transformed Oak Forests Humid Paramos Oak Forests Dry Paramos Transformed 851 34,375 2200 11,100 375 3800 119,150 16,200 1575 6050 1300 122,375 73,775 68,625 20,925 3775 10,675 20,800 5200 9475 129,375 22,000 62,275 4350 5675 27,125 8850 13,050 3125 3825 1150 7325 2550 2850 850 3575 % 100.0 71.5 4.6 23.1 0.8 2.7 84.7 11.5 1.1 82.3 17.7 40.8 24.6 22.9 7.0 1.3 3.6 58.6 14.7 26.7 59.3 10.1 28.6 2.0 10.4 49.6 16.2 23.9 38.6 47.2 14.2 74.2 25.8 39.2 11.7 49.1

PNN CORDILLER LOS PICACHOS

PNN CUEVA DE LOS GUACHAROS PNN EL COCUY

PNN PISBA

PNN SUMAPAZ

PNN TAMA

SFF GALERAS

PNN GUANENTA SFF IGUAQUE

Table 3 Fragmentation indices of the eastern Andes ecosystems, Colombiaa Ecosystem Dry forests Xeroc shrubs Subandean forests Andean forests Humid Paramos Dry Paramos Superparamo Nival Oak forests Intra andean savanahs Wetlands LPI 1.075 0.288 2.929 5.081 4.514 0.398 0.157 0.026 0.426 0.113 0.054 NP 135 16 302 118 40 6 3 4 19 4 7 MPS 2796.1 8043.7 5947.8 13,177.9 23,021.8 10,045.8 6975 943.7 6755.2 7487.5 2400 LSI 11.749 10.82 18.175 19.162 14.998 10.618 10.383 10.231 11.37 10.31 10.318 MNND 1446.4 15,444.4 1301.3 1602.7 6374.1 7859.6 4626.9 926.7 1566.8 3177 98,307.9

a LPI, largest patch index; NP, number of patches; MPS, mean patch size; LSI, landscape shape index; MNND, mean nearest neighbour distance.

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Andean and the sub Andean montane forests: 1.3 and 1.6 km. There were some dierences 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 transformed landscapes in which dispersed fragments of the original ecosystems occasionally appeared. The area between 4 and 8 300 N concentrated the greater number 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 geographically located around ve 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. inuence of Amazonian ecosystems and species) and high biological richness (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 represented in protected areas of the eastern Cordillera (35, 14.1 and 11%, respectively). This is due to the preponderance of highland parks in the Andes which originated in the establishment of Colombian protected natural areas without organized planning. However, when incorporating criteria based only on the percentage of total current area of an ecosystem, we are failing to consider the important reduction dating from preColumbian 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 gs. 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 dene whether or not an ecosystem is well protected within the national areas protected system, this is more a political conservation target than a scientically based result

and Sanjayan, 1998). Furthermore the declara(Soule tion of a protected area does not guarantee its protection, 7 of 11 national parks analyzed show some degree of transformation due to human activities and interactive management practices might be needed. Despite the high level of degradation, threat of habitat loss and further degradation and recognized scientic 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 identied as high priority region and the area for conservation: the Soacha-Bojaca around the protected area Estoraques (Fig. 4). Further work on a more detailed scale will allow better assessment of the real distribution of dry ecosystem remnants and the degree of anthropogenic disturbance. We suggest further protection of oak forest areas around the Sanctuaries of Flora and Fauna de Iguaque, and the -Alto Fonce river should also be considered Guanenta 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 identied 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 identied areas of interest are located in areas already within the national , Sumapaz and Picachos park system (Cocuy, Tama 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 geographic distribution and topographic landscape heterogeneity 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 specic soil and climatic conditions. Paramos and wetlands show similar restricted distributions. The mean neighbor distance and mean patch size of paramos was high because of its natural location in the highest mountains. Dry forest fragmented areas correspond to the Chicamocha canyon. Dry ecosystems have a high level of degradation and fragmentation that are not easy to dierentiate at this scale of research when observed from satellite images. These are only preliminary results that suggest conservation actions in specic 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

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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 specic information to identify a port-

folio of sites for conservation at dierent geographic levels. We also suggest redening some of the protected areas of the national parks system to include surrounding areas of natural ecosystems before they are further degraded, and in the mid term conduct an analysis of the

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

eective protection level of each area. There is also a need to clearly quantify land use change and deforestation rates through a multitemporal evaluation based on remote sensing images. This multitemporal evaluation of change will become a valuable criterion for dening 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

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their comments on the manuscript. This work was partially supported by The Nature Conservancy and Fun n Natura, Colombia. dacio

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