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ECOMOD-5004; No. of Pages 11 ARTICLE IN PRESS e c o l o g i c a l m o d e l l i n g x x x ( 2 0 0 7 ) xxx–xxx available at www.sciencedirect.com journal homepage: www.elsevier.com/locate/ecolmodel Spatially explicit models to analyze forest loss and fragmentation between 1976 and 2020 in southern Chile Cristian Echeverria a,b,∗ , David A. Coomes c , Myrna Hall d , Adrian C. Newton e a Departmento de Manejo de Bosques y Medio Ambiente, Facultad de Ciencias Forestales, Universidad de Concepción, Casilla 160-C, Concepción, Chile b Núcleo Cientı́fico Milenio FORECOS, Universidad Austral de Chile, Valdivia, Chile c Department of Plant Sciences, University of Cambridge, Cambridge CB2 3EA, United Kingdom d Faculty of Environmental Studies, SUNY College of Environmental Science and Forestry, 1 Forestry Drive, Syracuse, NY 13210, United States e School of Conservation Sciences, Bournemouth University, Talbot Campus, Poole, Dorset BH12 5BB, United Kingdom a r t i c l e i n f o a b s t r a c t Article history: Forest fragmentation threatens biodiversity in one of the last remaining temperate Received 9 June 2006 rainforests that occur in South America. We study the current and future impacts of frag- Received in revised form mentation on spatial configuration of forest habitats at the landscape level time in southern 8 October 2007 Chile. For this purpose, we identify the geophysical variables (“pattern drivers”) that explain Accepted 23 October 2007 the spatial patterns of forest loss and fragmentation between 1976 and 1999 using both a GIS-based land-use change model (GEOMOD) and spatially explicit logistic regression. Then, we project where and how much forest fragmentation will occur in the future by extrapola- Keywords: tion of the current rate of deforestation to 2010 and 2020. Both modeling approaches showed Deforestation consistent and complementary results in terms of the pattern drivers that were most related GEOMOD to deforestation. Between 1976 and 1999, forest fragmentation has occurred mainly from the Landscape indices edges of small fragments situated on gentle slopes (less than 10◦ ) and far away from rivers. Logistic regression We predict that patch density will decline from 2010 to 2020, and that total forest interior Temperate forests area and patch proximity will further decline as a result of forest fragmentation. Drivers Chile identified by these approaches suggest that deforestation is associated with observed local socio-economic activities such as clearance of forest for pasture and crops and forest logging for fuelwood. © 2007 Elsevier B.V. All rights reserved. 1. Introduction Fragmentation is a dynamic process in which the habitat of organisms is progressively reduced into smaller patches that become more isolated and affected by edge effects (Forman and Godron, 1986; Reed et al., 1996; Franklin, 2001). This process may lead to an increase in isolation of habitats and to a modification of ecosystem functioning that endangers species of plants, mammals and birds. Tropical forests have experienced a rapid, unprecedented forest loss and fragmentation that are having major impacts on wildlife, regional hydrology and the global climate (Laurance, 1999; Laurance et al., 2002). Similarly, temperate forests have been affected by land-cover change reaching high rates of forest loss in the southern hemisphere (Echeverrı́a et al., 2006). In particular, temperate rain forests in Chile, which are classified as a biological “hotspot” ∗ Corresponding autor at: Departmento de Manejo de Bosques y Medio Ambiente, Facultad de Ciencias Forestales, Universidad de Concepción, Casilla 160-C, Concepción, Chile. Tel.: +56 41 2204936; fax: +56 41 2255164. E-mail address: cristian.echeverria@udec.cl (C. Echeverria). 0304-3800/$ – see front matter © 2007 Elsevier B.V. All rights reserved. doi:10.1016/j.ecolmodel.2007.10.045 Please cite this article in press as: Echeverria, C., et al., Spatially explicit models to analyze forest loss and fragmentation between 1976 and 2020 in southern Chile, Ecol. Model. (2007), doi:10.1016/j.ecolmodel.2007.10.045 ECOMOD-5004; 2 No. of Pages 11 ARTICLE IN PRESS e c o l o g i c a l m o d e l l i n g x x x ( 2 0 0 7 ) xxx–xxx because of their high endemism (Myers et al., 2000), are disappearing as a result of land-cover change. This country has the largest temperate forest area in South America and more than a half of the total area of temperate forests in the southern hemisphere (Donoso, 1993). These forests are being harvested to supply the increasing global demand for wood and paper products and to clear areas for crops and pasture land (Lara et al., 2002). In response to growing concerns over the loss of biodiversity by fragmentation, specialists are seeking better ways of managing ecosystems at a variety of spatial and temporal scales (Turner et al., 2001). Growing evidence that habitat fragmentation may contribute substantially to the loss of regional or global biodiversity (Saunders et al., 1991) has provided empirical justification for the need to manage entire landscapes, not just the individual elements. For this purpose, landscape ecologists have developed techniques that provide in-depth analyses of the spatial configuration and compositional diversity of the elements of the landscape (McGarigal et al., 2002; Bennett, 2003; Herrmann et al., 2005). The development of geographical information system (GIS) techniques has offered a variety of analytical tools for the analysis at the landscape level. Many studies on deforestation and fragmentation analyze the temporal changes at a landscape level in order to understand the spatial patterns and interactions among the elements of the landscape and how these patterns change over time (Franklin, 2001; Staus et al., 2002; Fitzsimmons, 2003; Echeverrı́a et al., 2006). Also, ground-based studies provide data that can then be analyzed with respect to changing landscape patterns due to forest fragmentation (Metzger, 2000; Martı́nez-Morales, 2005; Echeverrı́a et al., 2007) in order to understand the consequences of these changes on composition and diversity of fragments (Cornelius et al., 2000; Asbjornsen et al., 2004; Castelletta et al., 2005). However, a more comprehensive understanding of the underlying drivers of fragmentation is required in order to inform future policy decision making or for land-use planning. This analysis must study the drivers responsible for forest loss and fragmentation, leading to the analysis of processes and not merely patterns (Bürgi et al., 2004). The use of the word “drivers” can be misleading since there are drivers or forcing functions that explain root causes and pressure on a system, and there are spatial pattern drivers, or landscape features, that influence where fragmentation is likely to occur. We focus on the latter, those geophysical attributes or landscape characteristics that determine where humans have chosen to invest their energies in order to survive (i.e. to derive food or income). These drivers show how the fragmentation has taken place spatially and temporally in the landscape. The ability to link a particular geophysical variable in the landscape to specific landscape changes is a powerful tool for researchers exploring environmental change (Evans and Moran, 2002). Modeling of land-cover changes such as deforestation requires combining spatially explicit ecological data with information on socio-economic factors (Dale and Pearson, 1997; Pearson et al., 1999). GEOMOD, a GIS-based model, can simulate the location of deforested cells using both biogeographical and socio-economic attributes as well as spatial data of forest cover at different time intervals (Hall et al., 1995a,b; Pontius et al., 2001). GEOMODs predictions can be improved by testing estimates of both the quantity of future forest areas as well as the location (Pontius, 2000). GEOMOD has the advantage that it does not require large amounts of data for calibration and validation, compared to other complex dynamic models (Pontius et al., 2001). Another advantage of GEOMOD over others is its use of validation prior to extrapolation. Logistic regression has been used to assess the probability of native forest conversion to industrial plantations in southern Chile (Wilson et al., 2005) and to create suitability maps of deforestation for the period 1971–1985 in Massachusetts, USA (Schneider and Pontius, 2001). Comparison of actual change maps to maps of modeled suitability revealed the ability of this approach to predict land-use changes. Serneels and Lambin (2001) used the logistic model to identify the drivers of conversion to agriculture in Narok District, Kenya, and Williams et al. (2005) applied it to determine the factor influencing the decline of native grasslands in Australia. Although analysis and modeling of deforestation are the focus of a substantial research effort of ecologists, very few studies have assessed the future impacts of fragmentation on spatial configuration of forest habitats at the landscape level by projecting the current trends of forest loss. Furthermore, relatively few studies have sought to compare different analytical approaches. The objectives of this study are (1) to identify the geophysical variables that determine forest cover change in southern Chile’s temperate forests using both GEOMOD and logistic regression; (2) to compare the results of the two analytical approaches; and (3) to assess the effects of such variables on the patterns of forest loss and fragmentation projected to the years 2010 and 2020. An understanding of landscape change is provided by integrating across time the causal relationships between forest cover change and spatially explicit variables. We hypothesize that changes in the configuration and area of the forest cover are the result of the preferred geophysical variables associated with forest logging and agriculture expansion. Also, we hypothesize that the current trends in deforestation may lead to substantial additional loss and fragmentation of the remaining forest fragments over the next decades. This information will be helpful to conservationist and biologists who are currently working on how to minimize the ecological impacts of ongoing deforestation. It will also contribute the provision of key information for decision making and management of natural resources. 2. Methods 2.1. Study area The study area covered approximately 500,000 ha located between 41◦ 30 –42◦ 20 S and 73◦ –74◦ W in southern Chile (Fig. 1). In the middle of the 20th century a portion of the native forests was cleared for agriculture as a result of European settlements (Donoso and Lara, 1997). The progressive deforestation in the landscape since 1985 may be related to a 400% increase in demand for industrial native forest products from 1988 to 1995 (Lara et al., 2002). This trend is explained mainly by the evolution of woodchip exports, which increased from 0.07 millions of tonnes in 1988 to 2.6 in 1995 (Lara et al., Please cite this article in press as: Echeverria, C., et al., Spatially explicit models to analyze forest loss and fragmentation between 1976 and 2020 in southern Chile, Ecol. Model. (2007), doi:10.1016/j.ecolmodel.2007.10.045 ECOMOD-5004; ARTICLE IN PRESS No. of Pages 11 e c o l o g i c a l m o d e l l i n g x x x ( 2 0 0 7 ) xxx–xxx 3 Fig. 1 – Location of the study area in southern Chile. 2002), and by a continuous production of fuelwood for growing municipalities located across the study landscape (Reyes, 2000). Expansion of crops and pastureland since 1970s has also been recognized as the other main cause of deforestation in the study area (Lara et al., 2002). In this process, recurrent fires are used as intermediate agent of conversion of forested areas to pasture. The study landscape is also highly representative of the region in terms of the forest composition. The native forests are characterized by the presence of several broad-leaved evergreen tree species such as Amomyrtus luma, Amomyrtus meli (both Myrtaceae), Drimys winteri (Winteraceae), Eucryphia cordifolia (Eucryphiaceae), Laurelia philippiana (Moniniaceae), Nothofagus dombeyi (Nothofagaceae), accompanied by a dense understory composed mainly of Chilean bamboos (Chusquea quila and Chusquea spp., Gramineae) and ferns. In some sites, the long-lived conifers such as Fitzroya cupressoides and Pilgerodendron uvifera (both Cupressaceae and highly threatened trees) can also be found. All these species grow both on deep soils of recent volcanic ash or on incompletely drained soil called Ñadi soil that present a solid iron-silicate-layer at a depth of 20–80 cm (Janssen et al., 2004). 2.2. Preparation of the spatial data To analyze change over a 23-year time period we acquired and classified a set of three Landsat satellite scenes: 1976 (MSS), 1985 (TM), and 1999 (ETM+). Each image was geometrically, atmospherically and topographically corrected. We used supervised classification and to assist in the classification of overlapping signatures, we applied the statistical decision criterion of maximum likelihood in which pixels were assigned to the class of highest probability (Chuvieco, 1996). The classification of the 1999 image was done using a set of thematic digital land-cover maps developed by one of the most comprehensive cartographic studies of natural vegetation conducted in Chile known as Catastro (CONAF et al., 1999), which covers the periods 1995–1997. A second reference group was comprised of 70 control points of field visits in which land-cover types that did not show changes in the last 30 years were recorded in July 2002. To carry out a quantitative comparison of the images, the original 79 m MSS raster grids were resampled to the resolution of the TM and ETM+ raster grids (30 m). The smallest patches (less than 5 pixels) were removed from all the images in order to reduce differences in data quality produced by the resampling of the MSS image. Images were classified into the following land-cover types: native forests, crop and pasture lands, shrublands and arboreus shrublands (both originated after forest logging or by regeneration in deforested areas), wetlands, water bodies and other land-cover types such as urban and bare areas. The overall accuracy of the classification of each image was estimated by constructing confusion matrices between reference data and classified data (Chuvieco, 1996). The accuracy of the ETM+ image was assessed by ground validation of 260 points. For the TM and MSS images, reference data (250 points for each image) was obtained from forest resource maps developed by the Instituto Forestal in 1979 and from additional field observations of land-cover types that did not exhibit changes between images over time. The overall accuracy values corresponded to 88.8% for the 1976 image, 89.6% for the 1985 image, and 91.9% for the 1999 image. The accuracy of native forests reached 94.9% in 1976, 95.9% in 1985 and 96.1% in 1999. The category with the lowest overall accuracy (67%) corresponded to the category of ‘other’ land-cover types in the 1976 image. The categories of land-cover types were grouped into the non-forest and forest categories to create a binary forest/nonforest map using ARC VIEW 3.2 software and its extension Arc View Spatial Analyst 2.0 for Window (ESRI, Redlands, CA, USA). In particular, crop and pasture lands, shrublands and Please cite this article in press as: Echeverria, C., et al., Spatially explicit models to analyze forest loss and fragmentation between 1976 and 2020 in southern Chile, Ecol. Model. (2007), doi:10.1016/j.ecolmodel.2007.10.045 ECOMOD-5004; No. of Pages 11 4 ARTICLE IN PRESS e c o l o g i c a l m o d e l l i n g x x x ( 2 0 0 7 ) xxx–xxx arboreus shrublands, wetlands and other land-cover types were grouped into the non-forest category, and native forests were grouped into the forest category. The maps were used to assess the patterns and drivers of forest loss and fragmentation. 2.3. Landscape spatial pattern analysis After reviewing recent forest fragmentation studies (Armenteras et al., 2003; Fitzsimmons, 2003; Imbernon and Branthomme, 2001; Millington et al., 2003; Staus et al., 2002), we selected from the landscape indices available in FRAGSTATS software (Version 3) (McGarigal et al., 2002) four landscape metrics that we believed would capture the resulting patterns of fragmentation over space and time. They included: (a) patch density (number of patches per 100 ha) as a measure of forest loss and division, (b) total edge length (km) as a measure of patch shape and the forest to clearance (cleared land) interface for potential edge effect studies, (c) total core area (total patch size in hectares) remaining after removing a buffer edge of 100 m; minimum distance defined according to previous studies that deal with edge effect (Millington et al., 2003; Vergara, 2005; Porej et al., 2004) as a measure of high-quality forest interior habitat, and (d) mean proximity index (ratio between the size and proximity of all patches whose edges are within a 1-km search radius of the focal patch) as a measure of isolation. These landscape indices were computed at the landscape level using the forest cover maps generated for each year. A Kruskal–Wallis test was used to determine if the indices obtained between time intervals were different at the 95% confidence level (Dytham, 2003). 2.4. Logistic regression We identified the areas deforested between each time interval (1976–1985 and 1985–1999) by overlaying the corresponding binary maps of forest/non-forest cover in a GIS (Fig. 2a). The binary response variable (forested, deforested) was analyzed using a logistic regression model. This model uses the following logit link function to transform the linear predictor (the combination of explanatory variables) to a measurement scale suitable for binomial data (Crawley, 2005): log p 1−p where p is probability. The probability of deforestation is expressed by the following model: p(x) = e(ˇ0 +ˇ1 X1 +···ˇk Xk ) 1 + e(ˇ0 +ˇ1 X1 +···ˇk Xk ) where Xk is the explanatory variable and ˇk is the estimated parameter of variable Xk . We selected a suite of nine candidate geophysical variables that we believe have influenced where people have cleared forest land for various economic uses over time (Fig. 2a). Most of these factors influence how much energy people will have to exert to get to the land and work the land and how much return they will experience from that investment. The variables selected include: slope, elevation, distance to roads in 1976, distance to rivers, distance to urban areas, soil type, dis- Fig. 2 – Flux diagrams illustrating the two spatially explicit modeling approaches used in this study: (a) logistic regression models and (b) GEOMOD modeling. tance to land already cleared for agricultural use in 1976, forest patch size, and distance to forest patch edge. We generated the first four variables from the Catastro data set at a scale of 1:50,000 (CONAF et al., 1999, Table 1). For soil types we used digital soil maps at a scale of 1:250,000 developed by Schlatter et al. (1995) and for maps of distance to 1976 agricultural area, patch size and distance to patch edge we used the land-cover maps derived from satellite images. We assumed that the contribution of these factors to deforestation has operated from 1976 to 1999 and will operate over the next decades. The geophysical and response variables were converted to a raster map of 30 m × 30 m grid cells using ©Arc View software (ESRI). Grid maps were exported into ©R 1.9.0 statistical software (Dalgaard, 2002) for logistic regression analyses. We randomly selected a sample of 1000 points for each time interval with the stipulation that they be separated by a distance of at least 1500 m. This distance lessened the effect of spatial autocorrelation. A proportion of 29.2% of the sampling points corresponded to deforested areas for the 1976–1986 period and 40%, for 1985–1999 period. We calculated their independence using the Moran index and achieved a value of 0.12, where Please cite this article in press as: Echeverria, C., et al., Spatially explicit models to analyze forest loss and fragmentation between 1976 and 2020 in southern Chile, Ecol. Model. (2007), doi:10.1016/j.ecolmodel.2007.10.045 ECOMOD-5004; No. of Pages 11 Geophysical variables GEOMOD modeling (1985–1999) Kappa (unconstrained) Kappa (constrained 3 × 3) Interval 1: 1976–1985 t-value Intercept Slope (◦ ) Distance to rivers (m) Elevation (m a.s.l.) Soil types Distance to roads (m) Distance to urban areas (km) Distance to 1976 agricultural areas (m) Patch size (ha) Distance to patch edge (m) 0–5, 5–10, 10–15, 15–20, 20–30, 30–40, 40–90 0–100, 100–200, 200–400, 400–800, 800–1,600, 1,600–3,200, 3,200–6,400, 6,400–12,800, >12,800 0–150, 150–300, 300–450, 450–600, >650 Volcanic ash, Ñadi, Marine sediments 0–300, 300–600, 600–1,200, 1,200–2,400, 2,400–4,800, >4,800 0–10, 10–20, 20–30, 30–40, >40 0–200, 400–600, 600–800, 800–1,000 >1,000 – – p-value Interval 2: 1985–1999 t-value p-value 1.01 −3.83 *** 0.5749 0.7426 0.72 −3.37 0.5841 0.7424 – n.s. 2.39 0.5909 0.7421 −0.86 n.s. −0.37 n.s. 0.5938 0.7417 – n.s. −0.17 n.s. 0.6381 0.7415 −1.69 n.s. −3.42 n.s. 0.5973 0.7414 0.56 n.s. 0.14 n.s. 0.5944 0.7404 – n.s. – n.s. – – – – n.s. *** −4.41 −4.11 – –8.01 *** * *** *** N = 1000 points; *p < 0.01; ***p < 0.001. ARTICLE IN PRESS Categories Logistic regression modeling e c o l o g i c a l m o d e l l i n g x x x ( 2 0 0 7 ) xxx–xxx 5 Please cite this article in press as: Echeverria, C., et al., Spatially explicit models to analyze forest loss and fragmentation between 1976 and 2020 in southern Chile, Ecol. Model. (2007), doi:10.1016/j.ecolmodel.2007.10.045 Table 1 – Attribute maps and/or explanatory variable incorporated in GEOMOD simulations and logistic regression models, respectively. Attribute maps were used in categories in GEOMOD and as continuous explanatory variables (except for soil types) in logistics regressions. The highest kappa (0.7430) value was obtained by combining slope and distance to rivers into the simulations. Values of kappa were obtained for simulations constrained and unconstrained by the nearest neighbor search mode. Variables without t-values correspond to those variables that were not entered in the multivariate logistic model because of their no significance in their respective univariate models ECOMOD-5004; No. of Pages 11 6 ARTICLE IN PRESS e c o l o g i c a l m o d e l l i n g x x x ( 2 0 0 7 ) xxx–xxx 0 indicates no spatial dependence and 1.0 indicates a high degree of spatial autocorrelation. Variables weakly correlated (r < 0.5) were selected to fit the logistic models. For each time interval, we separately entered each geophysical variable into ©S-PLUS 6.0 statistical software (Crawley, 2005) to generate a univariate logistic regression model (Fig. 2a). A variable was retained if the change in deviance between the model containing the variable and the null model was significant at a p-value <0.05. Then, the significant geophysical variables were entered into the model simultaneously to generate a multivariate logistic regression model for each time interval (Fig. 2a). The function “drop1” of R was used to assess the significance of the variables in each model at a p-value <0.05. Drop1 is a backward selection method available in R that tests the change in deviance against a 2 distribution. 3. GEOMOD modeling GEOMOD selects locations where changes in land uses are most likely by using three decision rules based on: (a) the pattern of geophysical variables with respect to already deforested land, (b) stratification by political sub-region, and/or (c) nearest neighbors. This rule simulates the manner in which deforestation occurs on the edge or in open areas of forest fragments. For the GEOMOD analysis we used the same set of geophysical variables tested in the logistic regression analysis (Table 1), except patch size and distance to patch edge that cannot be entered as mapped variables for GEOMOD analysis. However, distance to patch edge is captured by GEOMODs nearest neighbor rule, which constrains the selection of cells to those that lie along the border of forest patches. GEOMOD constructed a suitability map (areas more vulnerable to be deforested) based on the degree to which the geophysical variables have determined past deforestation patterns (Fig. 2b). The suitability map was created in two steps (Pontius et al., 2001). First, attribute maps were binned into three to nine categories (Table 1). Second, the model reclassified each grid cell of each category of the geophysical variable by assigning a percent-disturbed value, obtained by comparing the geophysical map to the initial land-use map. GEOMOD computed the percent-disturbed value of each category (bin) in the geophysical variable which is the ratio of the quantity of disturbed grid cells of that category to all cells of that category in the map. The final suitability map for each model run is the weighted sum of all reclassified attribute maps to be used in that run. Hence, we calculated the suitability in each cell according to the following: R(i) = A a=1 Wa Pa (i) where R(i) is the suitability value in cell (i), a the particular geophysical variable map, A the number of geophysical variable maps, Wa the weight of the geophysical variable map a, and Pa (i) the percent-disturbed in category ak of the geophysical variable a, where cell (i) is a member of category ak . To avoid assigning an arbitrary importance to the variables, simulations were run using a weight (W) equal to 1 for each variable. The method of calibration consisted in deriving a simulated map for 1999, using the 1985 map as a base map and then simulating forest loss at the rate recorded between 1985 and 1999 (Fig. 2b). The iterative process of calibration and validation to find those variables that yielded the best-fit model consisted of comparing the simulated 1999 map to the actual map for the simulated year using the kappa-for-location parameter (Pontius, 2002) (Fig. 2b). This parameter validates the simulation’s ability to predict location and is equal to 1 when simulation’s success rate is perfect and is equal to 0 when the simulation’s success rate is equivalent to that due to chance (Pontius, 2000). In our case, a “success” occurred when the grid cell in the map of simulated land-use matched the corresponding grid cell in the actual map in 1999. The effects of GEOMODs decision rules (nearest neighbor and geophysical variable weighting) were examined using each variable individually with and without the nearest neighbor mode. The selection of the most important spatial pattern drivers to be used in the forward projection to 2010 and 2020 was based on those variables that achieved the highest values of kappa-for-location upon validation against the 1999 map (Fig. 2b). After selecting the “best-fit” geophysical variables, we ran simulations of forest cover to 2010 and 2020. The quantity of future forest cover was determined by linearly extrapolating the forest loss rate observed in the classified imagery between 1985 and 1999 which amounts to 1691 ha year−1 equivalent to 0.78% year−1 . It was assumed that both the rate and the influence of geophysical variables would remain constant over time. The extrapolated 2010 and 2020 forest cover maps used for the evaluation of the four landscape indices corresponded to maps of gross deforestation, as our analysis assumes that there is no regrowth of deforested areas. 4. Results 4.1. Logistic regression models The multivariate logistic regression model proposed that the probability of an area being cleared of forest for the 1976–1985 interval was highly significant (p < 0.001) and negatively related to distance from patch edges and slope (Table 1). The variables considered in the logistic model of deforestation after exclusion of highly correlated variables were patch size, slope, elevation, soil type, distance to rivers, distance to patch edge, and distance to towns. In the univariate logistic regression five variables explained some of the variation in the pattern of forest cover at p < 0.001, except patch size (2 = 1.8, d.f. = 1, p = 0.179), distance from river (2 = 2, d.f. = 1, p = 0.157), distance to 1976 agricultural areas (2 = 1.1, d.f. = 1, p = 0.219), and soil types (F = 0.095, d.f. = 1, p = 0.922), which did not show a significant change in deviance. Therefore, these variables were not entered in the backward selection (no t-value in Table 1). For the 1985–1999 interval, the probability of deforestation was related negatively to distance from patch edges, slope and patch size and positively to distance from rivers (Table 1). During this period, all the explanatory variables were significant, except distance from 1976 agricultural areas in 1976 (Table 1). The most powerful predictors were patch size and distance to patch edge. Please cite this article in press as: Echeverria, C., et al., Spatially explicit models to analyze forest loss and fragmentation between 1976 and 2020 in southern Chile, Ecol. Model. (2007), doi:10.1016/j.ecolmodel.2007.10.045 ECOMOD-5004; No. of Pages 11 ARTICLE IN PRESS e c o l o g i c a l m o d e l l i n g x x x ( 2 0 0 7 ) xxx–xxx 4.2. GEOMOD modeling Simulations revealed that the most effective decision rule to model the location of forested cells was the one based on slope, distance to rivers and the application of a 3 × 3 nearest neighbor deforestation rule (Table 1). In this case, GEOMOD was able to classify correctly 88.04% of the grid cells (kappa = 0.7430) of the 1999 reference map. Higher kappa values (>0.74) were obtained when the simulations in GEOMOD (from 1985 to 1999) were constrained to the nearest neighbor search mode (Table 1). Individually slope yielded the highest kappa with 0.7426, followed by distance to rivers with 0.7424. The categories of slope with ranges 0–5◦ presented the highest percent of deforested cells in 1985 with 58%. The lowest percent-deforested (7%) was concentrated in slopes between 20◦ and 30◦ . For areas situated between 200 and 800 m away from rivers, the percent-deforested was 60%. In forests located between 0 and 100 m from rivers almost 50% of cells were deforested. With respect to elevation, the majority of deforested cells (58%) were concentrated between 0 and 150 m a.s.l. Only 8% of deforested cells occurred higher than 450 m a.s.l. Although the predictive power of soil type was lower (kappa = 0.7417), almost 60% of cells situated in volcanic ash soil were deforested, whereas 45% of deforestation occurred in Ñadi soil. At less than 300 m from the nearest road, 80% of the cells were deforested. For a distance greater than 4.8 km, this percent declined to 22%. Forests at less than 20 km 7 from urban areas were 54% deforested. More than 80% of the cells located less than 800 m from areas under cultivation in 1976 were deforested by 1985. 4.3. Extrapolation of forest cover In 1985, 46% of the study area was forested, but in 1999 this decreased to 41% (Fig. 3). In our extrapolations, the native forests of the study area were reduced to 38% by 2010 and 35% by 2020 (Fig. 3). This represents a loss of approximately 66,700 ha of native forest in 35 years. Although our analysis assumes that there is no regrowth of deforested areas (as described in Section 2), there is reforestation evidence in the satellite imagery of approximately 7% between 1976 and 1985 and 11% between 1985 and 1999. 4.4. Observed and future trends in forest fragmentation For the years observed in our analysis, patch density increased from 1.41 patches/100 ha in 1976 to 2.5 in 1999 (Fig. 4a). From 1999 to 2010, this index did not exhibit changes. However, during the last time interval, the patch density declined to 2.2 patches/100 ha. These modifications in the configuration of the forest cover were also associated with greater patch edge length between forest and non-forest cover types (Fig. 4b). The overall variation in these values indicated that the shape Fig. 3 – Spatial and temporal patterns of forest cover in southern Chile for the years 1976, 1985, 1999, 2010, and 2020. Forest cover maps for the years 2010 and 2020 are based on simulations that apply the annual deforestation rate between 1985 and 1999, geophysical attribute maps of slope and distance to rivers, and application of the 3 × 3 nearest neighbor rule in GEOMOD. Please cite this article in press as: Echeverria, C., et al., Spatially explicit models to analyze forest loss and fragmentation between 1976 and 2020 in southern Chile, Ecol. Model. (2007), doi:10.1016/j.ecolmodel.2007.10.045 ECOMOD-5004; No. of Pages 11 8 ARTICLE IN PRESS e c o l o g i c a l m o d e l l i n g x x x ( 2 0 0 7 ) xxx–xxx Fig. 4 – Changes in landscape indices applied to forest cover from 1976 to 2020. Values of indices derived from actual maps (1976, 1985, and 1999) are depicted in solid black lines. Black-dashed lines represent extrapolated values based on the forest loss rate between 1985 and 1999 (0.78% year−1 ). Landscape indices: (a) patch density (number of patches per 100 ha), (b) total edge length, (c) total core area (for a distance to edge of 100 m), and (d) mean proximity index (for a distance of 1 km). of forest patches became more irregular up to 1999. In our simulated results total edge length of forest patches in 2020 decreased significantly (12 = 66.42, p < 0.001) (Fig. 4b) over that of 2010. From 1976 the total core area constantly decreased up to 1999, but for the extrapolated years, the total core area did not appear to be significantly reduced between 1999 and 2010 (12 = 1.33, p = 0.25) and between 2010 and 2020 (12 = 0.22, p = 0.63) (Fig. 4c). The mean proximity index, or measure of isolation, sharply declined up to 1999. Although the index recorded in 2010 was relatively similar to that one recorded in 1999, a significant reduction of this index was found (12 = 2322.4, p < 0.001). Similarly, the index of proximity continued to significantly decline by 2020 (12 = 156.05, p < 0.001) (Fig. 4d). 5. Discussion 5.1. Modeling approaches GEOMOD was applied to identify the major spatial attributes that drive landscape change as well as to simulate the progressive loss of native forest into the future. GEOMOD was able to classify correctly 88% of grid cells of the simulated 1999 map, recording a kappa of 0.74, which is categorized as “very good” by Monserud and Leemans (1992). A slightly higher kappa of 0.80 was obtained using GEOMOD for land-cover change analysis in northeastern Massachusets (Pontius et al., 2003). A strong similarity was recorded here between the two modeling approaches used in identifying the geophysical variables associated with forest loss. The univariate logis- tic regression demonstrated that each explanatory variable, excluding distance to 1976 agricultural land, yielded a high probability of deforestation between 1985 and 1999. The importance of these variables as drivers of deforestation was confirmed by GEOMOD, which showed high values of kappa for each variable, with the lowest kappa achieved using distance to 1976 agricultural land as the independent driver. For the same period, the multivariate logistic regression revealed that the most significant explanatory variables (excluding patch size and distance to patch edge that were not analyzed in GEOMOD) corresponded to those with the highest values of kappa obtained by GEOMOD, i.e. slope and distance to rivers. The use of logistic regression models enabled GEOMODs results to be supplemented. For the two time intervals, the logistic models determined that distance to patch edges was highly significant in explaining the probability of deforestation. This is consistent with the analysis conducted using GEOMOD, in which the simulations that produced the highest kappa were done applying the neighborhood search mode of 3 × 3 cells. In this mode, GEOMOD constrains the selection of cells to be deforested to those cells that are around the border of forest patches. Some researchers state that the difficulty in predicting the location of deforested areas is related to the fact that they are scattered evenly with respect to underlying factors and some of the possibly most important variables are not available in digital format (Schneider and Pontius, 2001). In the current study, the deforestation processes have been concentrated in certain sites in the landscape, which enabled significant geophysical variables to be identified. Even though most of the variables used in this study come from the most detailed and Please cite this article in press as: Echeverria, C., et al., Spatially explicit models to analyze forest loss and fragmentation between 1976 and 2020 in southern Chile, Ecol. Model. (2007), doi:10.1016/j.ecolmodel.2007.10.045 ECOMOD-5004; No. of Pages 11 ARTICLE IN PRESS e c o l o g i c a l m o d e l l i n g x x x ( 2 0 0 7 ) xxx–xxx 9 comprehensive survey available for Chile (CONAF et al., 1999), a number of different scales have been combined in this analysis, so that the detail required to understand a process at the highest map scale is missing. A limitation of the study is related to the fact that the predicted forest maps are based on the assumption that the rate of deforestation between 1985 and 1999 will remain constant over the next decades. Wilson et al. (2005) used logistic regression to assess the probability of native forest conversion to industrial plantations in southern Chile. That study was limited to the assumption that plantations were converted directly from native forests, excluding the possibility that plantations could have been established in previously deforested areas such as crops, shrublands, and grasslands. In the present work, the availability of forest covers generated at three different time intervals enabled more complex patterns of land-use conversion to be distinguished. For each time interval, we were able to identify those cells that were previously occupied by forests and then converted to different land-cover types. flood risk. These restrictions were more evident in flat areas where the forest patches were intensely fragmented and left as riparian vegetation. Similar results were found in Melbourne, Australia, where the application of logistic regression to analyze the decline of native grassland revealed that patches close to streams presented a low probability of destruction (Williams et al., 2005). Forested flat areas near towns and roads were highly vulnerable to native forest replacement by plantations of exotic species in southern Chile (Wilson et al., 2005). Similarly, the proximity to village settlements was found to be among the most important variable of forest conversion to agricultural land in Madagascar (McConnell et al., 2004). In contrast to these studies, we found that distance to towns and distance to roads were not significant in accounting for clearance of forest area for the two time intervals examined. However, the high values of kappa obtained by GEOMOD for these geophysical variables show that these attributes are also important to be able to predict the location of forested areas by 1999. 5.2. Relating our findings to what we observe on the ground 5.3. Forest loss up until 1976 was most highly correlated to the presence of volcanic ash soil, which is located in lowlands and is the most favorable for vegetation growth (INIA, 1985), such as agricultural crops and cattle grazing. Through our analysis we have found that between 1976 and 1999 the expansion of crops and pasture lands continued to occur in Ñadi soil, which is very limited in soil depth, and in gently sloping terrain where more and more forested areas were opened for grazing and cultivation. This process has resulted in recurrent fires, which act as an intermediate agent of conversion from forest to pasture lands. In both time periods analyzed in the present study, the clearance of forests was concentrated around edges of forest fragments located in slightly undulating terrain. This pattern is related to the fact that local people use trees near the borders of forest patches to produce fuelwood and then clear these areas for crops and pasture land. Likewise, a logistic model-based study conducted in Madagascar also found that the expansion of agriculture into the remaining natural forest was associated with progressive clearance from forest edges (McConnell et al., 2004). In our study area, the spatial pattern of forest logging during the observed years was the result of an increase in the demand for native forest products such as firewood and woodchips during the 1980s (Lara et al., 2002). Currently, forest logging for firewood is the main factor driving forest loss in the study area (Lara et al., 2002). We found that during the first time interval, patch size was not a significant variable due to the presence of forest fragments of different sizes that occurred in flat areas. However, between 1985 and 1999, small patches became vulnerable to deforestation, because they were mainly concentrated in flat areas, where it is easier to clear forest land, thus leading to intensification of fragmentation in these areas. Additionally, pattern analysis revealed that the clearance of forests took place mainly in patches located away from rivers or streams. This finding correlates with the fact that clearance of forests is legally prohibited in areas close to rivers or in areas with Future trends in forest fragmentation Based on the current trends of the geophysical variables driving deforestation, a continuous reduction in the area of intermediate and large fragments is expected during the next two decades. Due to that there is evidence of reforestation between 1976 and 1999, it is probable that the predicted areas of native forests in 2010 and 2020 are slightly higher than our estimates. Because forests are continuously being affected by fragmentation over time, a decline in patch density is observed from 2010 to 2020. This index increased in the earliest stage of forest loss and fragmentation and then decreases during the extrapolated years. This means that if the current trends continue, the process of deforestation may even eliminate forest patches created in the last decades. The identification of this threshold in the patch density has also been reported further north in central Chile, where the decline in patch density was recorded approximately 20 years ago due to rapid deforestation (Echeverrı́a et al., 2006). Similarly, Zipperer et al. (1990) observed that the constant action of deforestation led to a decline in patch density in central New York, USA. The native forests of our study area presented abrupt changes in their spatial configuration over the whole study period, from a forest habitat formed by complex clusters of large fragments to a sparse distribution of smaller patches. This trend of decreasing connectivity of forest fragments is the result of increasing forest fragmentation through time, which has dissected the native forests into more compact, isolated, and smaller fragments. The isolation of fragments is a fundamental consequence of unchecked fragmentation and may have negative impacts if the populations of species occurring in these habitats have no capacity to survive in isolated fragments or to move through the surrounding modified matrix (Willson et al., 1994; Cornelius et al., 2000; Reed and Levine, 2005; Echeverrı́a et al., 2007). Results revealed a slight decrease of the core area and proximity of forest fragments in the next two decades. This is not the result of a decline in the forest loss rate, but a change in Please cite this article in press as: Echeverria, C., et al., Spatially explicit models to analyze forest loss and fragmentation between 1976 and 2020 in southern Chile, Ecol. Model. (2007), doi:10.1016/j.ecolmodel.2007.10.045 ECOMOD-5004; No. of Pages 11 10 ARTICLE IN PRESS e c o l o g i c a l m o d e l l i n g x x x ( 2 0 0 7 ) xxx–xxx the spatial configuration in which most remaining forests are becoming more inaccessible or restricted owing to legal regulations on forest logging in steep areas and next to streams. The study landscape in 2020 is characterized by a mosaic of small fragments, probably influenced by edge effects, highly isolated from large intact forests located at higher elevations. Similarly, future projections by Pearson et al. (1999) in the temperate forest of the USA, produced complex changes in the spatial patterns, by increasing the number of patches and decreasing the size of some habitats suitable for the species studied. 6. Conclusions The results of our analysis demonstrate the advantages of using standard remote sensing data along with spatially explicit modeling approaches to identify and assess the major geophysical variables that influence the pattern and location of deforestation and fragmentation. Results demonstrated that the pattern of deforestation and fragmentation may have a notable effect on the spatial configuration of the remaining forest fragments over the next decades and potential detrimental consequences on biodiversity conservation. The spatial patterns described in this research reveal the immediate drivers of deforestation and fragmentation that operate in the study area. These drivers suggest that forest logging and clearance for crops and pasture land are the main human activities associated with the changes in the spatial configuration of forest cover. Future studies that assess the relationship between immediate drivers and underlying causes of deforestation such as environmental legislation, political economy, poverty, etc., should be conducted for a more comprehensive understanding of the causes of forest loss in the world. Acknowledgements This research was made possible by funding from BIOCORES (biodiversity conservation, restoration, and sustainable use in fragmented forest landscapes) project ICA-CT-200110095; bursary for young researchers from developing countries—European Union (administered by UFZ, Germany); the UNEP-WCMC Chevening Scholarship in biodiversity; the Frank Smart Studentship of the Department of Plant Sciences, University of Cambridge; the support from Cambridge Commonwealth Trust & Cambridge Overseas Trust 2003-2005; FORECOS (P04-065-F) Millennium Scientific Nucleus of MIDEPLAN; and ALFA-FOREST (contract II-0411-FA-FCD-FI-FC). CE thanks Patricio Romero for his assistance in the preparation of digital covers. references Armenteras, D., Gast, F., Villareal, H., 2003. Andean forest fragmentation and the representativeness of protected natural areas in the eastern Andes, Colombia. Biol. Conserv. 113, 245–256. Asbjornsen, H., Ashton, M., Vogt, D., Palacios, S., 2004. Effects of habitat fragmentation on the buffering capacity of edge environments in a seasonally dry tropical oak forest ecosystem in Oaxaca, Mexico. Agric. Ecosyst. Environ. 103, 481–495. Bennett, A., 2003. Linkages in the Landscape. The Role of Corridors and Connectivity in Wildlife Conservation. IUCN, Gland, Switzerland and Cambridge, UK, 254 pp. Bürgi, M., Hersperger, A.M., Schneeberger, N., 2004. Driving forces of landscape change—current and new directions. Landsc. Ecol. 19, 857–868. Castelletta, M., Thiollay, J., Sodhi, N., 2005. The effects of extreme forest fragmentation on the bird community of Singapore Island. Biol. Conserv. 121, 135–155. Chuvieco, E., 1996. Fundamentos de teledetección espacial. Ediciones RIALP, S.A., Third ed., Madrid, Spain, 568 pp. CONAF, CONAMA, BIRF, Universidad Austral de Chile, Pontificia Universidad Católica de Chile, Universidad Católica de Temuco, 1999. Catastro y Evaluación de los Recursos Vegetacionales Nativos de Chile. Informe Nacional con Variables Ambientales. Santiago, Chile, 88 pp. Cornelius, C., Cofre, H., Marquet, P., 2000. Effects of habitat fragmentation on bird species in a relict temperate forest in semiarid Chile. Conserv. Biol. 14, 534–543. Crawley, M., 2005. Statistical Computing. An Introduction to Data Analysis using S-PLUS. Wiley, USA, 761 pp. Dale, V.H., Pearson, S.M., 1997. Quantifying habitat fragmentation due to land use change in Amazonia. In: Laurance, W., Bierregaard, R. (Eds.), Tropical Forest Remnants: Ecology, Management, and Conservation of Fragmented Communities. The University of Chicago Press, Chicago, USA, pp. 400–414. Dalgaard, P., 2002. Introductory Statistics with R. Springer, 267 pp. Donoso, C., 1993. Bosques templados de Chile y Argentina: variación, estructura y dinámica. Universitaria, Santiago, Chile, 484 pp. Donoso, C., Lara, A., 1997. Utilización de los bosques nativos en Chile: pasado, presente y futuro. In: Armesto, J.J., Villagrán, C., Arroyo, M.K. (Eds.), Ecologı́a de los bosques nativos de Chile. Editorial Universitaria, Santiago, Chile, pp. 335–362. Dytham, C., 2003. Choosing and using Statistics: A Biologist’s Guide. Blackwell Science, York, UK, pp. 218. Echeverrı́a, C., Coomes, D., Salas, J., Rey-Benayas, J.M., Lara, A., Newton, A., 2006. Rapid deforestation and fragmentation of Chilean temperate forests. Biol. Conserv. 130, 481–494. Echeverrı́a, C., Coomes, D., Newton, A., Rey-Benayas, J.M., Lara, A., 2007. Impacts of forest fragmentation on species composition and forest structure in the temperate landscape in southern Chile. Global Ecol. Biogeogr. 16, 426–439. Evans, T., Moran, E., 2002. Spatial integration of social and biophysical factors related to land cover change. Popul. Dev. Rev. 28, 165–186. Fitzsimmons, M., 2003. Effects of deforestation and reforestation on landscape spatial structure in boreal Saskatchewan, Canada. Forest Ecol. Manage. 174, 577–592. Forman, R.T.T., Godron, M., 1986. Landscape Ecology. John Wiley and Sons, USA, 619 pp. Franklin, S., 2001. Remote Sensing for Sustainable Forest Management. Lewis Publishers, USA, 407 pp. Hall, C.A.S., Tian, H., Qi Y., Pontius, G., Cornell, J., Uhlig, J., 1995a. Spatially explicit models of land use change and their application to the tropics. DOE Research Summary, No. 31. CDIAC, Oak Ridge National Lab. Hall, C.A.S., Tian, H., Qi, Y., Pontius, G., Cornell, J., Uhlig, J., 1995b. Modeling spatial and temporal patterns of tropical land use change. J. Biogeogr. 22, 753–757. Herrmann, H., Babbitt, K., Baber, M., Gongalton, R., 2005. Effects of landscape characteristics on amphibian distribution in a Please cite this article in press as: Echeverria, C., et al., Spatially explicit models to analyze forest loss and fragmentation between 1976 and 2020 in southern Chile, Ecol. Model. (2007), doi:10.1016/j.ecolmodel.2007.10.045 ECOMOD-5004; No. of Pages 11 ARTICLE IN PRESS e c o l o g i c a l m o d e l l i n g x x x ( 2 0 0 7 ) xxx–xxx forest-dominated landscape. Biol. Conserv. 123, 139–149. Imbernon, J., Branthomme, A., 2001. Characterization of landscape patterns of deforestation in tropical rain forests. Inter. J. Remote Sens. 22, 1753–1765. Instituto de Investigaciones Agropecuarias (INIA), 1985. Suelos volcánicos de Chile. Ministerio de Agricultura, Santiago, Chile. Janssen, I., Kruemmelbein, J., Hora, R., Ellies, A., 2004. Physical and hydraulic properties of the Nadi soils in south Chile—comparison between untilled and tilled soil. Revista de la Ciencia del Suelo y Nutrición Vegetal 4, 14–28. Lara, A., Echeverrı́a, C., Reyes, R., 2002. Bosques. In: Centro de Análisis de Polı́ticas Públicas. Universidad de Chile (Ed.). Informe Paı́s: Estado del Medio Ambiente en Chile-2002. Santiago, Chile, pp. 131–160. Laurance, W.F., 1999. Reflections on the tropical deforestation crisis. Biol. Conserv. 91, 109–117. Laurance, W.F., Lovejoy, T.E., Vasconcelos, H.L., Bruna, E.M., Didham, R.K., Stouffer, P.C., Gascon, C., Bierregaard, R.O., Laurance, S.G., Sampiao, E., 2002. Ecosystem decay of Amazonian forest fragments: a 22-year investigation. Conserv. Biol. 16, 605–618. Martı́nez-Morales, M., 2005. Landscape patterns influencing bird assemblages in a fragmented neotropical cloud forest. Biol. Conserv. 121, 117–126. McConnell, W., Sweeney, S., Mulley, B., 2004. Physical and social access to land: spatio-temporal patterns of agricultural expansion in Madagascar. Agriculture. Ecosyst. Environ. 101, 171–184. Metzger, J.P., 2000. Tree functional group richness and landscape structure in a Brazilian tropical fragmented landscape. Ecol. Applic. 10, 1147–1161. McGarigal, K., Cushman, S.A., Neel, M.C., Ene, E., 2002. Fragstats: spatial pattern analysis program for categorical maps. Retrieved January 20, 2003, from University of Massachusetts, Landscape Ecology Program web site: http://www.umass.edu/landeco/research/fragstats/fragstats.html. Millington, A.C., Velez-Liendo, X.M., Bradley, A.V., 2003. Scale dependence in multitemporal mapping of forest fragmentation in Bolivia: implications for explaining temporal trends in landscape ecology and applications to biodiversity conservation. J. Photogr. & Remote Sens. 57, 289–299. Monserud, R.A., Leemans, R., 1992. Comparing global vegetation maps with the Kappa statistic. Ecol. Model. 62, 275–293. Myers, N., Mittermeler, R.A., Mittermeler, C.G., da Fonseca, G.A.B., Kent, J., 2000. Biodiversity hotspots for conservation priorities. Nature 403, 853–858. Pearson, S., Turner, M., Drake, J., 1999. Landscape change and habitat availability in the southern Appalachian Highlands and Olympic Peninsula. Ecol. Applic. 9, 1288–1304. Pontius, R.G., 2000. Quantification error versus location error in comparison of categorical maps. Photogr. Eng. Remote Sens. 66, 1011–1016. Pontius, R.G., 2002. Statistical methods to partition effects of quantity and location during comparison of categorical maps at multiple resolutions. Photogr. Eng. Remote Sens. 68, 1041–1049. 11 Pontius, R.G., Cornell, J., Hall, C., 2001. Modelling the spatial pattern of land-use change with GEOMOD2: application and validation for Costa Rica. Agric. Ecosyst. Environ. 85, 191–203. Pontius, R.G., Agrawal, A., Huffaker, D., 2003. Estimating the uncertainty of land-cover extrapolations while constructing a raster map from tabular data. J. Geogr. Syst. 5, 253–273. Porej, D., Micacchion, M., Hetherington, T.E., 2004. Core terrestrial habitat for conservation of local populations of salamanders and wood frogs in agricultural landscapes. Biol. Conserv. 120, 399–409. Reed, R., Johnson-Barnard, J., Baker, W., 1996. Fragmentation of a forested rocky mountain landscape, 1950–1993. Biol. Conserv. 75, 267–277. Reed, M., Levine, S., 2005. A model for behavioral regulation of metapopulation dynamics. Ecol. Model. 183, 411–423. Reyes, R., 2000. Caracterización de los sistemas de producción y comercialización de leña para la ciudad de Puerto Montt, X Región. Thesis, Facultad de Ciencias Forestales, Universidad de Chile, Santiago, Chile. Saunders, D.A., Hobbs, R.J., Margules, C.R., 1991. Biological consequences of ecosystem, fragmentation: a review. Conserv. Biol. 5, 18–32. Schlatter, J., Gerding, V., Huber, H., 1995. Sistema de Ordenamiento de la Tierra. Herramienta para la planificación forestal aplicado a la X Región. Serie Técnica. Facultad de Ciencias Forestales. Universidad Austral de Chile, Valdivia. Schneider, L., Pontius, R.G., 2001. Modeling land-use change in the Ipswich watershed, Massachusetts, USA. Agric. Ecosyst. Environ. 85, 83–94. Serneels, S., Lambin, E., 2001. Proximate causes of land-cover change in Narok District, Kenya: a spatial statistical model. Agric. Ecosyst. Environ. 85, 65–81. Staus, N., Strittholt, J., Dellasala, D., Robinson, R., 2002. Rate and patterns of forest disturbance in the Klamath-Siskiyou ecoregion, USA between 1972 and 1992. Landsc. Ecol. 17, 455–470. Turner, M.G., Gardner, R.H., O’Neill, R.V., 2001. Landscape Ecology in Theory and Practice: Pattern and Process. Springer, New York, 497 pp. Vergara, P., 2005. Predation on bird nests in forest edges of a National Reserve of the Coastal Range of the Maule Region. In: Smith-Ramirez, C., Armesto, J.J., Valdovinos, C. (Eds.), Historia, biodiversidad y ecologiı̌a de los bosques costeros de Chile. Editorial Universitaria, Santiago, Chile, pp. 578–585. Williams, N.G., McDonnell, M.J., Seager, E., 2005. Factors influencing the loss of an endangered ecosystem in an urbanizing landscape: a case study of native grasslands from Melbourne, Australia. Landsc. Urban Plan. 71, 35–49. Willson, M., De Santo, T.I., Sabag, C., Armesto, J.J., 1994. Avian communities of fragmented south-temperate rainforests in Chile. Conserv. Biol. 8, 508–520. Wilson, K., Newton, A., Echeverrı́a, C., Weston, C., Burgman, M., 2005. A vulnerability analysis of the temperate forests of south central Chile. Biol. Conserv. 122, 9–21. Zipperer, W.C., Burgess, R.L., Nyland, R.D., 1990. Patterns of deforestation and reforestation in different landscape types in central New York. Forest Ecol. Manage. 36, 103–117. Please cite this article in press as: Echeverria, C., et al., Spatially explicit models to analyze forest loss and fragmentation between 1976 and 2020 in southern Chile, Ecol. Model. (2007), doi:10.1016/j.ecolmodel.2007.10.045