Impacts of Anthropogenic Land-Use Change on Populations of The

Impacts of anthropogenic land-use change on populations of the Endangered Patagonian cypress Fitzroya cupressoides in southern Chile: implications for its conservation

J AMES R ODRÍGUEZ-ECHEVERRY,CRISTIAN E CHEVERRÍA and L AURA N AHUELHUAL

Abstract Land-use change can have negative effects on Introduction threatened species by modifying their habitat and population dynamics. The habitat of the Endangered and-use change can lead to habitat loss and fragmen- Patagonian cypress Fitzroya cupressoides (Mol.) Johnst. Ltation, and these are two of the greatest threats to forest   (Cupressaceae), a conifer endemic to the temperate forests ecosystems (Noss, ; Baillie et al., ). Land-use of southern Chile and Argentina, has been transformed by change modifies connectivity, and patch density and area  land-use change and overexploitation. The impact of land- of forest (Lindenmayer & Fischer, ). These changes use change on the spatial pattern of F. cupressoides habitat can influence ecological attributes at the species and com- from  to  was evaluated at the landscape level, using munity levels, such as species richness and individual den-   satellite images. Additionally, eight  ×  m plots were es- sity (Laurance et al., ; Bustamante et al., ;  tablished in four populations to assess their status. In each Simonetti et al., ). Habitat loss and fragmentation can ’ plot the density of F. cupressoides and the species richness of affect species survival by reducing the quality and quantity  the associated plant communities were recorded and ana- of habitat available (Tomimatsu & Ohara, ), both of ’ lysed, together with spatial patterns at the population and which can alter the dynamics of a species population. community levels. The loss of potential F. cupressoides habi- Such alteration could lead to local and global extinction of  tat was % from  to  (–% for the four popu- the most vulnerable species (Bennett, ). lations). The density of F. cupressoides was lowest in smaller The impact of land-use change on threatened species  habitat patches, where the number of plant species was warrants investigation (Turner, ; Lindenmayer &  higher and the matrix dominated by anthropogenic land- Fischer, ) to facilitate understanding of the way in use. Land-use change was associated with a loss of potential which it can alter the composition and spatial configuration habitat for F. cupressoides, and differences in the spatial pat- of the unique habitats on which threatened species depend, terns of habitat influenced the composition of remaining the influence of such alterations on species composition, the populations and communities. A landscape approach is re- influence of the matrix, and the minimum buffer zone commended as a conservation planning strategy for F. needed for management of the remaining interior habitats  cupressoides. (Oliveira et al., ). Multi-scale studies (landscape, habitat and species) could provide valuable information for the Keywords Anthropogenic processes, Chilean larch tree, conservation of threatened species (Vergara & Simonetti, Fitzroya cupressoides, habitat assessment, landscape change, ). Patagonian cypress, spatial patterns Chilean temperate forest, which has been designated a To view supplementary material for this article, please visit hotspot for biodiversity conservation because of its high  http://dx.doi.org/S level of endemism (Myers et al., ), has experienced significant degradation, fragmentation and loss since  as a result of land-use change (Echeverría et al., ). Consequently there have been significant changes in richness of plant (Echeverría et al., ) and bird species (Vergara & Simonetti, ) and the spatial distribution  JAMES RODRÍGUEZ-ECHEVERRY (Corresponding author) and CRISTIAN ECHEVERRÍA of threatened species (Altamirano et al., ). Departamento de Manejo de Bosques y Medio Ambiente, Facultad de The Patagonian cypress Fitzroya cupressoides, an en- Ciencias Forestales, Universidad de Concepción, Chile E-mail [email protected] demic and monotypic conifer of the temperate forests of southern Chile and Argentina, is categorized as LAURA NAHUELHUAL Facultad de Ciencias Agrarias, Instituto de Economía  Agraria, Universidad Austral de Chile, Chile Endangered on the IUCN Red List (Premoli et al., ). .  Received  May . Revision requested  September . Individuals can live for , years, making it the second Accepted  October . First published online  March . longest-lived species and thus of high scientific value (Lara

Oryx, 2015, 49(3), 447–452 © 2015 Fauna & Flora International doi:10.1017/S0030605314000945 Downloaded from https://www.cambridge.org/core. IP address: 170.106.202.8, on 25 Sep 2021 at 05:22:25, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0030605314000945 448 J. Rodríguez-Echeverry et al.

& Villalba, ). Since the arrival of Europeans in the south use was categorized as native forest, shrubland, wetland, of Chile in  F. cupressoides has been overexploited for its grassland, forest plantations of exotic species or other uses beautiful and decay-resistant wood. In Chile the species is (bare soil, urban areas and water bodies). Changes in spatial found in the Coastal Range, the Andean Range and the patterns of the forest landscape were assessed by determin- Central Depression (Fraver et al., ). The species is ing the total area and number of patches of native forest, known from  sites in the Central Depression, five contain- using FRAGSTATS v. . (McGarigal et al., ) and the ing small remnant populations and eight supporting scat- Spatial Analyst extension for ArcGIS v. .. (ESRI, tered trees and saplings in a landscape transformed by Redlands, USA). human activities such as logging for firewood and expansion Four of the five known F. cupressoides populations in the of pastureland for cattle grazing (Fraver et al., ). Genetic Central Depression were included in the study; the fifth studies indicate that the F. cupressoides populations in the population was not assessed because access to information Central Depression are the most genetically distinct of the about it was limited. Spatially explicit information on the lo- remaining populations in Chile and Argentina (Premoli cations of populations was provided by Corporación et al., ), and represent ancient populations from the gla- Nacional Forestal. cial era, from which the other populations are derived The habitat of F. cupressoides in the Central (Premoli et al., ). Depression is characterized by poorly drained acidic Studies have focused on the reproductive biology, ecol- soils (pH of .)knownasñadis (Fig. ;FAO- ogy and distribution of F. cupressoides (Armesto et al., UNESCO, ), which means seasonal swamp in the in- ; Donoso et al., ; Fraver et al., ) but none have digenous Mapudungun language. These soils have a quantified the impact of land-use change on the spatial pat- depth of – cm, abundant humus, moss beds and terns of the species’ habitat and examined the effect of habi- coarse woody debris. The habitat of F. cupressoides is tat fragmentation and loss on population composition. This also characterized by a cold oceanic temperate climate, type of information is necessary to assess the current status with annual mean temperature and rainfall of .°C of F. cupressoides populations, which may have been signifi- and , mm, respectively, and by the presence of the fol- cantly altered in Chile. lowing species of plants: Amomyrtus luma, Drimys win- We assessed the impact of land-use change on the spatial teri, Laureliopsis philippiana, Saxegothaea conspicua and pattern of the forest landscape and the habitat of F. cupres- Weinmannia trichosperma (IUCN, ). soides in the Central Depression in southern Chile, and A previous study in the same area revealed a relationship related changes in spatial pattern to the composition of between F. cupressoides habitat and the interior of small four remaining populations. We hypothesize that human- patches of native forest (Fraver et al., ), which had a induced land-use change has substantially fragmented core area of , ha and a buffer zone of $ m. Fraver the remaining F. cupressoides habitat, resulting in re- et al. () reported that in the patches of native forest duced population density and variation in community where edge effects (altered species composition, community composition. structure and microclimate) occurred within  m of the patch edge, F. cupressoides populations were not present and only a few scattered F. cupressoides were recorded.  Study area Following the approach of Fraver et al. ( ) we defined the potential habitat of F. cupressoides as an area of native  The study area (, km ) was defined by the distribution of forest , ha with a buffer zone of $ m. four remaining populations of F. cupressoides in the Central Areas of potential habitat for F. cupressoides were ident- Depression, Chile (Fraver et al., ), and by Landsat satel- ified by analysing the core area and buffer zone of native for- lite images used to determine land-use change (Fig. ). The est patches, using the Land Change Modeler for Ecological area, characterized by a cold oceanic climate, has a maxi- Sustainability extension of IDRISI Andes (Clark Labs, mum elevation of  m, mean temperature of .°C and Worcester, USA), which generated maps of potential habitat annual rainfall of , mm. The landscape in this area is for the years  and . Changes in spatial patterns of dominated by patches of broad-leaved evergreen native for- potential habitat were evaluated by analysing the total area est of the Valdivian Rainforest. (ha) and number of potential habitat patches over time, using FRAGSTATS. Subsequently we identified the habitat patches that include the four remaining populations of Methods F. cupressoides. In  we recorded the tree density (based on individuals Land-use change analyses were performed based on maps of with diameter at breast height (DBH) $ cm and height land use, with  x  m pixels, derived from Landsat satel- $ m) and level of regeneration (based on seedlings with lite images for  and  (Echeverría et al., ). Land DBH , cm and/or height , m) in the four F. cupressoides

FIG. 1 Location of the four surveyed populations (–)ofFitzroya cupressoides in the Central Depression, Chile. The rectangle on the inset shows the location of the main map in Chile.

populations (Fig. ). We established two  x  m sampling to , ha. In  the four F. cupressoides populations plots randomly within each forest patch, taking into account occurred in patches of – ha. In  these patches were the small size of the patches and the likelihood of edge ef- .– ha (Table ). Losses of , ,  and % of potential fects. To facilitate data collection each plot was divided habitat were recorded over the entire study period for popu- into   x  m subplots, following the methods of Peet lations , ,  and , respectively (Table ). et al. (). We estimated the number of plant species pres- There appeared to be a correlation between lower density ent in native forest patches where F. cupressoides was pres- of F. cupressoides seedlings and trees and greater loss of po- ent (Jiménez-Valverde & Hortal, ). This information tential habitat (Table ). In  for the population with a was analysed together with the area of the native forest loss of % of potential habitat we recorded a density of patches where F. cupressoides was present, and the area of , trees per ha and regeneration of , trees per ha. F. cupressoides habitat and the matrix that surrounded For the population with a loss of % of potential habitat each F. cupressoides population. the density was  trees per ha and there was no regeneration (Table ). The populations of F. cupressoides for which greater loss Results of habitat was recorded were those surrounded by an anthropogenic matrix and were associated with plant com- There was a loss of % of native forest in the study land- munities with the greatest number of native and exotic spe- scape from  to  (i.e. a mean annual loss of .%; cies (Table ). In  the population for which a % loss of Supplementary Fig. S). In  there were , patches potential habitat was recorded was located in the largest of native forest, with a total area of , ha. The number patch of native forest ( ha), embedded in a matrix domi- of patches in  (,) was more than double that in nated by native forest, and associated with a community , with a reduction in total area to , ha, and shrub- with the lowest richness of exotic plant species (Table ). land had become the dominant land cover type The population for which a % loss of potential habitat There was a % loss in potential habitat for F. cupres- was recorded was located in the smallest patch of native for- soides from  to  (i.e. a mean annual loss of .%). est (. ha), embedded in a matrix dominated by grassland, In  potential habitat for the species occurred in  and associated with a community with the greatest richness patches of native forest (a total of , ha). In  the of exotic plant species (Plantago truncata, Poa annua and number of patches had decreased to  and the total area Prunella vulgaris; Table ).

Discussion . Changes in the forest landscape and Fitzroya  cupressoides habitat

Our results indicate there was a substantial loss of native for- No. of exotic species in sampling plots est in the study landscape from  to , although the   was present in annual rate of loss ( . %) was lower than that recorded for other hotspot landscapes that have been significantly transformed, such as the Maulino temperate forest in central Chile (.% annual loss; Echeverría et al., ) and the tropical montane forest of Chiapas, Mexico (.% annual F. cupressoides  Species richness of associated plant communities No. of native species in sampling plots loss; Cayuela et al., ). The loss of native forest area

) was associated with an increase in the number of patches, 1 − from , in  to , in , which is one of the main symptoms of fragmentation (Lindenmayer & Fischer, ). The trend in fragmentation is similar to

F. cupressoides that reported for other temperate landscapes in Chile Density of seedlings (ha where threatened tree species occur (Bustamante & ) 1 

− Castor, ), as well as in tropical montane landscapes of conservation importance in Mexico (Cayuela et al., ). This trend of loss and fragmentation of native forest in Composition of populations Density of trees (ha the study landscape was associated with a significant loss (%) of potential habitat for F. cupressoides since the – early s, with a decrease in the number of patches of potential habitat from  to . If the current trajectory of loss shrubland 26 2,270 4,340 7 1 62 40 0 18 3 36 360 2,470 9 2 46 340and 1,350 fragmentation 16 of native forest 3 continues, further losses – – – – – of potential habitat for F. cupressoides can be expected. 24 36 29 51 – – – – Forest grassland matrix (%) Changes at the population and community level in 2011

We observed differences in habitat loss in each of the four F. potential cupressoides populations, associated with the densities of the populations and number of plant species in patches of native forest where populations. The smallest habitat patches had the lowest densities of F. cupressoides. Similar trends have been reported for other threatened plant species in Chile, such as F. cupressoides Nothofagus alessandrii (Bustamante & Castor, ) and Legrandia concinna (Altamirano et al., ), for various Loss of habitat since 1999 (%) species of birds (Vergara & Simonetti, ; Simonetti

Fitzroya cupressoides et al., ), and for some populations of insectivorous birds, primates and mammals in the Amazon rainforest (Laurance et al., ). A reduction in habitat size may decrease the density of F. cupressoides populations and thus increase the risk of extinction for this species. Our results show that differences in the spatial patterns potential habitat (ha) of habitat for the four F. cupressoides populations were associated with differences in the plant species richness of the associated communities. As the size of F. cupressoides habitat patches decreased, the number of native and exotic plant Area of forest patches with F. cupressoides Habitat spatial patterns species increased. In the Chilean temperate forest a similar trend in community composition was reported for plants  1 Habitat spatial patterns, composition of (Bustamante & Grez, ), birds (Vergara & Simonetti, ) and small mammals (Kelt, ). In boreal (Chávez ABLE Population 127 38 50 2 0.4 100 2 312 54 35 4 0.5 79 3 T & Macdonald, ) and tropical areas (Trauernicht &

Ticktin, ) this change is less dramatic, perhaps because planning focused on sustainable production practices such forest landscapes have not been transformed to the same ex- as agroforestry. The design and implementation of such a tent. Other factors may also influence changes in the plant strategy requires identification and evaluation of sensitive species richness of communities associated with patches of buffer areas and their connectivity. Moreover, this strategy native forest where F. cupressoides occurs. The matrix may must be supported by a framework of environmental induce several types of abiotic changes at the edge of forest policies. patches, resulting in the potential establishment and recruit- We recommend that the Chilean government expands ment of generalist plant species (Bustamante et al., ). the country’s protected areas to conserve F. cupressoides po- The combination of different patch sizes and matrix compo- pulations and promote conservation of the species. At pres- sition may have influenced the establishment of generalist ent there is only one protected F. cupressoides population in native and exotic species in each community where F. cu- the Central Depression, which is the Monumento Natural pressoides was present. Lahuen Ñadi. We also recommend strengthening the exist- ing restoration programmes for F. cupressoides, and de- Implications for conservation veloping new programmes, to improve the quality of habitats for F. cupressoides. These programmes could in- The Central Depression is located in a landscape that has clude patches of native forest that are potential habitat for been subjected to constant anthropogenic pressure the species, monitoring to assess the viability and success (Torrejón et al., ). From the th to the th century of conservation efforts, and investment of capital. significant changes in the spatial patterns of native forest Researchers from the Universidad Austral de Chile, land were caused mainly by extensive wood extraction owners, and Corporación Nacional Forestal are already par- (Torrejón et al., ), whereas since the sthegreatest ticipating in restoration programmes (Premoli et al., ) changes in the spatial patterns of native forests have been but the conservation of F. cupressoides also requires the caused by commercial plantations and expansion of pas- commitment and active participation of the Chilean govern- tureland for cattle grazing (Echeverría et al., , ). ment and the general community. The landscape of the Central Depression is under increasing pressure from the cities of Puerto Montt and Puerto Varas (Fraver et al., ), and anthropogenic land use Acknowledgements may result in further changes in the spatial patterns of na-  tive forest and F. cupressoides habitat (Armesto et al., ; We thank FONDECYT project , CONAF, Alberto Wilson et al., ). Hernández, Natalia Medina, Víctor Sandoval, and Fitzroya cupressoides was listed in Appendix I of CITES Laboratorio de Ecología de Paisaje of the Universidad de in  and declared a Natural Monument by the Chilean Concepción. We also thank the Ministry of Education of government in . Our multi-scale study has shown a pro- Chile for a MECESUP scholarship. gressive and severe loss of native forest with F. cupressoides populations from  to . If habitat loss continues, References the population density of F. cupressoides may decrease

further, increasing the risk of extinction for the species. As ALTAMIRANO, A., ECHEVERRÍA,C.&LARA,A.() Efecto de la a first step in conserving F. cupressoides we suggest a land- fragmentación forestal sobre la estructura vegetacional de las scape approach, using the corridor–patch–matrix model poblaciones amenazadas de Legrandia concinna (Myrtaceae) del  – (Lindenmayer & Franklin, ), as a strategy for conser- centro-sur de Chile. Revista Chilena de Historia Natural, , . ARMESTO, J., SMITH-RAMIREZ, C., LEÓN,P.&ARROYO,M.() vation planning. This is appropriate given the current con- Biodiversidad y conservación del bosque templado en Chile. figuration of the landscape, with native forest restricted to Ambiente y Desarrollo, , –. small patches sparsely distributed across the landscape. BAILLIE, J.E.M., HILTON-TAYLOR,C.&STUART, S.N. ()  The main objective of the corridor–patch–matrix model is IUCN Red List of Threatened Species: A Global Species Assessment. to maintain the quality and quantity of patches of native for- IUCN, Gland, Switzerland.  est through management of the matrix (Lindenmayer & BENNETT, A.F. ( ) Linkages in the Landscape: The Role of Corridors  and Connectivity in Wildlife Conservation. IUCN, Gland, Franklin, ). The condition of the matrix may be more Switzerland, and Cambridge, UK. important in determining the survival of the species than BUSTAMANTE, R.O. & CASTOR,C.() The decline of an endangered the isolation of patches (Lindenmayer & Franklin, ). temperate ecosystem: the ruil (Nothofagus alessandrii) forest in In the study landscape the management of the matrix central Chile. Biodiversity & Conservation, , –.  should focus on sensitive buffer areas that improve the con- BUSTAMANTE, R.O. & GREZ, A.A. ( ) Consecuencias ecológicas de la fragmentación de los bosques nativos. Ambiente y Desarrollo, , nectivity between forest patches and increase the ability of –. the matrix to support F. cupressoides populations. We rec- BUSTAMANTE, R.O., SEREY, I.A. & PICKETT, S.T.A. () Forest ommend that this strategy be complemented with land-use fragmentation, plant regeneration and invasion processes across

edges in central Chile. In How Landscapes Change: Human Maps. Http://www.umass.edu/landeco/research/fragstats/fragstats. Disturbance and Ecosystem Fragmentation in the Americas html [accessed  September ]. (eds G.A. Bradshaw & P.A. Marquet), pp. –. Springer-Verlag, MYERS, N., MITTERMEIER, R.A., MITTERMEIER, C.G., DA FONSECA, Berlin, Germany. G.A.B. & KENT,J.() Biodiversity hotspots for conservation CAYUELA, L., REY BENAYAS, J.M. & ECHEVERRÍA,C.() Clearance priorities. Nature, , –. and fragmentation of tropical montane forests in the Highlands of NOSS, R.F. () Forest fragmentation in the Southern Rocky Chiapas, Mexico (–). Forest Ecology and Management, , Mountains. Landscape Ecology, , –. –. OLIVEIRA, M.A., GRILLO, A.S. & TABARELLI,M.() Forest edge in CHÁVEZ,V.&MACDONALD, S.E. () The influence of canopy the Brazilian Atlantic forest: drastic changes in tree species patch mosaics on understory plant community composition in assemblages. Oryx, , –. boreal mixedwood forest. Forest Ecology and Management, , PEET, R.K., WENTWORTH, T.R., DUNCAN,R.&WHITE, P.S. () –. North Carolina Vegetation Survey Protocol: A Flexible, Multipurpose CITES () Convention on International Trade in Endangered Method for Recording Vegetation Composition and Structure Species of Wild Fauna and Flora. Appendices I, II and III. Http:// (version .). University of North Carolina, Chapel Hill, USA. cites.org/eng/app/appendices.php [accessed  December ]. PREMOLI, A., QUIROGA, P., SOUTO,C.&GARDNER,M.() Fitzroya DONOSO, C., SANDOVAL, V., GREZ,R.&RODRÍGUEZ,J.() cupressoides.InIUCN Red List of Threatened Species v. .. Http:// Dynamics of Fitzroya cupressoides forests in southern Chile. Journal www.iucnredlist.org [accessed  December ]. of Vegetation Science, , –. PREMOLI, A., VERGARA, R., SOUTO, C., LARA,A.&NEWTON,A. ECHEVERRÍA, C., COOMES, D., SALAS, J.M., REY BENAYAS, J., LARA,A. () Lowland valleys shelter the ancient conifer Fitzroya &NEWTON,A.() Rapid deforestation and fragmentation of cupressoides in the Central Depression of southern Chile. Journal of Chilean temperate forests. Biological Conservation, , –. the Royal Society of New Zealand, , –. ECHEVERRÍA, C., NEWTON, A., LARA, A., REY BENAYAS, J.M. & SIMONETTI, J.A., GREZ, A.A. & BUSTAMANTE, R.O. () COOMES, D.A. () Impacts of forest fragmentation on species Interacciones y procesos en el bosque maulino fragmentado. In composition and forest structure in the temperate landscape of Biodiversidad en ambientes fragmentados de Chile: patrones y southern Chile. Global Ecology and Biogeography, , –. procesos a diferentes escalas (eds A.A. Grez, J.A. Simonetti & R. ECHEVERRÍA, C., NEWTON, A., NAHUELHUAL, L., COOMES,D.&REY O. Bustamante), pp. –. Editorial Universitaria, Santiago, Chile. BENAYAS, J.M. () How landscapes change: integration of spatial TOMIMATSU,H.&OHARA,M.() Genetic diversity and local patterns and human processes in temperate landscapes of southern population structure of fragmented populations of Trillium Chile. Applied Geography, , –. camschatcense (Trilliaceae). Biological Conservation, , –. FAO-UNESCO () Soil Map of the World. Volume IV: South TORREJÓN, F., CISTERNAS, M., ALVIAL,I.&TORRES,L.() America. UNESCO, Paris, France. Consecuencias de la tala maderera colonial en los bosques de alerce FRAVER, S., GONZÁLEZ, M.E., SILLA, F., LARA,A.&GARDNER,M. de Chiloé, sur de Chile (siglos XVI–XIX). Magallania, , –. () Composition and structure of remnant Fitzroya cupressoides TRAUERNICHT,C.&TICKTIN,T.() The effects of non-timber forests of southern Chile’s Central Depression. Journal of the Torrey forest product cultivation on the plant community structure and Botanical Society, , –. composition of a humid tropical forest in southern Mexico. Forest IUCN () IUCN Red List of Threatened Species v. .. Http:// Ecology and Management, , –. www.iucnredlist.org [accessed  December ]. TURNER, M.G. () Landscape ecology: the effect of pattern on JIMÉNEZ-VALVERDE,A.&HORTAL,J.() Las curvas de process. Annual Review of Ecology and Systematics, , –. acumulación de especies y la necesidad de evaluar la calidad VERGARA, P.M. & SIMONETTI, J.A. () Avian responses to de los inventarios biológicos. Revista Ibérica de Aracnología, fragmentation of the Maulino Forest in central Chile. Oryx, , , –. –. KELT, D.A. () Small mammal communities in rainforest WILSON, K., NEWTON, A., ECHEVERRÍA, C., WESTON,C.& fragments in Central Southern Chile. Biological Conservation, , BURGMAN,M.() A vulnerability analysis of the temperate –. forests of south central Chile. Biological Conservation, , –. LARA,A.&VILLALBA,R.()A-year temperature record from Fitzroya cupressoides tree rings in southern South America. Science, Biographical sketches , –. LAURANCE, W.F., VASCONCELOS, H.L. & LOVEJOY, T.E. () Forest JAMES RODRÍGUEZ-ECHEVERRY is a biologist interested in loss and fragmentation in the Amazon: implications for wildlife landscape ecology, ecological restoration and conservation. His re- conservation. Oryx, , –. search focuses on global change, threatened species, biodiversity and LINDENMAYER, D.B. & FISCHER,J.() Habitat Fragmentation and ecosystem services in the landscapes of Latin America. CRISTIAN Landscape Change. An Ecological and Conservation Synthesis. Island ECHEVERRÍA is a forester interested in landscape ecology and conser- Press, Washington, DC, USA. vation. His research focuses on threatened species, landscape change LINDENMAYER, D.B. & FRANKLIN, J.F. () Conserving Forest and ecological restoration of degraded forests in Latin America. Biodiversity: A Comprehensive Multiscaled Approach. Island Press, LAURA NAHUELHUAL is an agronomist interested in the economics Washington, DC, USA. of natural resources and rural development. Her research focuses on MCGARIGAL, K., CUSHMAN, S.A., NEEL, M.C. & ENE,E.() assessment and economic valuation of ecosystem services in Latin FRAGSTATS: Spatial Pattern Analysis Program for Categorical America.