Latawiec Et Al 2015

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See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/276473638

Creating space for large-scale restoration intropical agricultural landscapes

ARTICLE in FRONTIERS IN ECOLOGY AND THE ENVIRONMENT MAY 2015

Impact Factor: 7.44 DOI: 10.1890/140052

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Bernardo Strassburg

Pontifcia Universidade Catlica do Rio de Ja

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Pedro H. S. Brancalion

Escola Superior de Agricultura "Luiz de Queir

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Ricardo R. Rodrigues

University of So Paulo

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Toby Alan Gardner

Stockholm Environment Institute

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The Ecological Society of America www.frontiersinecology.org

The anthropogenic degradation of natural ecosystems,including land-use conversion, has led to widespreadloss of biodiversity and has compromised the provision ofecosystem services in landscapes around the world(Cardinale et al. 2012). There are now approximately twobillion hectares of deforested and degraded land world-wide (Minnemeyer et al. 2011). As a consequence ofefforts to reverse this trend, ecological restoration is nowwidely recognized as a global environmental priority

(Aronson and Alexander 2013). The Aichi Target 15 ofthe United Nations Convention on Biological Diversity,an international instrument for the conservation and sus-tainable use of biodiversity, together with the BonnChallenge a global restoration initiative have estab-lished a goal of restoring 150 million hectares of defor-ested and degraded land globally by 2020. The New YorkDeclaration on Forests expands this goal to 350 millionhectares restored by 2030. In addition, several large-scalerestoration initiatives have emerged around the world inrecent years for instance, the Green Belt Movement inKenya (de Aquino et al. 2011), where millions of hectaresof agricultural land are expected to be restored soon(Roberts et al. 2009; Rodrigues et al. 2011).

In many parts of the world, the restoration of ecosystemservices and the conservation of biodiversity depend onthe success of such projects. Even if only partly successful,such initiatives could become yet another major competi-tor for land an increasingly scarce resource that isneeded to satisfy the growing demands of a progressivelylarger and more affluent human population (Smith et al.2010). Consequently, a key challenge in the managementof any large-scale restoration project is avoiding the dis-

placement of pre-existing land uses such as agriculture, asthis may result in further clearance of native vegetationand loss of biodiversity elsewhere (Meyfroidt and Lambin2009; Melo et al. 2013). This displacement could alsoproduce negative social consequences for the people whohave been displaced (Barr and Sayer 2012).

There is evidence that such displacement, or leakage,has already occurred in some locations. For example,about 39% of the increase in forest cover in Vietnambetween 1987 and 2006 appears to have been balanced bycommensurate increases in deforestation in nearby coun-tries (eg Laos, Cambodia, and Indonesia; Meyfroidt and

CONCEPTS AND QUESTIONS

Creating space for large-scale restoration intropical agricultural landscapes

Agnieszka E Latawiec1,2,3,4, Bernardo BN Strassburg1,4*, Pedro HS Brancalion5, Ricardo R Rodrigues6,and Toby Gardner1,7

Poorly planned, large-scale ecological restoration projects may displace agricultural activities and potentially

lead to the clearance of native vegetation elsewhere, with associated impacts on biodiversity and ecosystem ser-

vices. Yet few studies have considered these risks and the ways in which restoration can increase competition for

land. Here, we address this issue by examining whether large-scale restoration of the Brazilian Atlantic Forest

could displace cattle production, as a result of land shortages. Although the risks of displacement are indeed

high when reforestation is planned in areas with high cattle productivity, we discuss how these risks can be

minimized through a combination of productivity increases, a regional restoration planning framework, and

the prioritization of marginal agricultural land for restoration. We also consider how restoration can, in some

circumstances, be made more economically sustainable by incorporating income-generating activities such as

exploitation of timber and non-timber forest products, certification, and payments for ecosystem services.

Front Ecol Environ 2015; 13(4): 211218, doi:10.1890/140052

1International Institute for Sustainability, Rio de Janeiro, Brazil*([email protected]); 2Department of Production Engineering

and Logistics, Opole University of Technology, Opole, Poland; 3School

of Environmental Science, University of East Anglia, Norwich, UK;4Sustainability Lab, Department of Geography and the Environment,

Pontificia Universidade Catolica, Rio de Janeiro, Brazil; 5Department

of Forest Sciences, University of So Paulo, Piracicaba, Brazil;6Department of Biology, University of So Paulo, Piracicaba, Brazil;7Stockholm Environment Centre, Stockholm, Sweden

In a nutshell: Large-scale ecological restoration can displace pre-existing

agricultural activities and may drive increased competition forland, resulting in the loss of native vegetation elsewhere

It is possible to reconcile large-scale restoration and agricul-tural expansion through improvements in cattle ranching pro-ductivity in regions with intense competition for land, such asthe Atlantic Forest of Brazil

The farming sector, agricultural institutions, and planningauthorities all have a central role to play in any large-scalerestoration program, helping to avoid negative socio-environ-mental outcomes as a result of poorly planned restorationwhile promoting compliance with environmental laws

Further research is urgently needed to study and monitor the

effects of large-scale restoration on people and the environment


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Large-scale restoration in tropical landscapes AE Latawiec et

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www.frontiersinecology.org The Ecological Society of Amer

Lambin 2009). This pattern of displaced land use is alsoevident in other countries where large-scale restorationoccurred (Meyfroidt et al. 2010). Yet while the threat ofindirect effects of land-use change has received increas-ing attention in the context of agricultural expansion (egsoybeans and sugarcane in Brazil, Arima et al. 2011; cross-biome leakage worldwide, Strassburg et al. 2014a), it hasreceived unexpectedly little attention in the context oflarge-scale ecological restoration. In fact, many legiti-mate concerns about potential negative consequences of

increased competition for land from restoration projectsare largely based on anecdotal evidence (Barr and Sayer2012) and very little spatially explicit information isavailable on the potential for conflicts over land betweenagriculture and restoration.

Here, we examine the potential for future competitionfor land between forest restoration and cattle pastures inBrazils Atlantic Forest biome and estimate the potentialfor displacement of cattle pastures. We also consider howany such displacement could be avoided or mitigated atboth local and national levels. In particular, we assessalternative ways of reconciling multiple demands for land

through increases in the productivity of cattle ranchinbased on a case study of the Brazilian state of EspriSanto. To this we apply alternative scenarios assessinexisting state-imposed targets for the expansion of agrculture, plantation forestry, and forest restoration.

The Atlantic Forest is one of the most species-rich bimes on the planet, yet less than 15% of the original fore

is left; much of what remains is highly degraded, makinthis region a national and global priority for restoratioefforts. The case study offered by the Atlantic Forest wfacilitate exploring the consequences of increased comptition for land as a result of large-scale restoration, boin Brazil and across the tropics, and may help in develoing a conceptual foundation to promote further advancin research and policy.

Opportunities for restoration in low-productivity

pasturelands

The Atlantic Forest is a unique and highly threaten

biome, retaining only 814% of its original areal cover 150 million hectares; of the remaining forest fragments, 80are less than 50 ha in size (Figure 1; Ribeiro et al. 2009Even though it is also one of the top five global biodiversihotspots, referred to often as the hottest of the hotspot(Laurance 2009), it still experiences annual deforestatiorates of more than 20000 ha (Soares-Filho et al. 2014). addition, the Atlantic Forest biome is of great importanfor Brazilian society, providing a home for approximate60% of the national population (IBGE 2012); about 80%the countrys gross domestic product is generated within thAtlantic Forest boundaries (IBGE 2012).

The Atlantic Forest Restoration Pact (AFRP; a grouof environmental organizations, private companies, goernment agencies, researchers, and landowners) in Brais a biome-scale restoration initiative aimed at maximing the benefits of large-scale forest restoration whiavoiding negative environmental and social outcom(Melo et al. 2013). The target areas for restoration weselected by the AFRP, based on an assessment of defoested lands included within so-called PermanePreservation Areas (APP in Portuguese) in accordanwith the Brazilian Forest Code; totaling seven milliohectares, these areas primarily include riparian buffzones along streams and around springs (where restor

tion is mandatory), as well as extensive, low-productivipasturelands with few cattle (Calmon et al. 2011Together, these lands total 17 728 187 ha, providing tbasis for the AFRPs stated goal of restoring 15 milliohectares of forest within the Atlantic Forest biome b2050 (Calmon et al. 2011; Melo et al. 2013). There acurrently approximately 30 million hectares of plantpastures in the Atlantic Forest, supporting 36 milliohead of cattle at an average stocking rate of 0.82 animunits (AU) per hectare (1 head equals 0.7 AU; PROBI2009) a very low level of production efficiency by intenational standards (Strassburg et al. 2014b).

Figure 1. The Atlantic Forest, a global biodiversity hotspot

despite being highly fragmented. Approximately 30 million hectaresin the Atlantic Forest region are used for cattle ranching.

MR

angel/InternationalInstituteforSustainability
https://www.researchgate.net/publication/248200076_Conserving_the_hottest_of_the_hotspots_Preface?el=1_x_8&enrichId=rgreq-1775e3f5-c006-4ab0-b1ab-ea7318af2cd4&enrichSource=Y292ZXJQYWdlOzI3NjQ3MzYzODtBUzoyODA0NjE2MjU1Nzc0NzNAMTQ0Mzg3ODY3MDMwMg==https://www.researchgate.net/publication/248200076_Conserving_the_hottest_of_the_hotspots_Preface?el=1_x_8&enrichId=rgreq-1775e3f5-c006-4ab0-b1ab-ea7318af2cd4&enrichSource=Y292ZXJQYWdlOzI3NjQ3MzYzODtBUzoyODA0NjE2MjU1Nzc0NzNAMTQ0Mzg3ODY3MDMwMg==


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AE Latawiec et al. Large-scale restoration in tropical landscapes

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Restoration success will presumably vary considerablyacross the Atlantic Forest biome for several reasons,including variability in both natural soil conditions(Sobanski and Marques 2014) and historical human use.Restoration is likely to be more effective (ie supportingthe long-term recovery of a species-rich, functioning for-est ecosystem) and to require fewer expensive interven-

tions (eg soil preparation, active plantings, clearance ofexotic trees and grasses to prevent competitive exclusion)in areas that have been subject to less intense and exten-sive levels of historical use and in areas that are closer toforest remnants, which provide source populations ofnative species. Given these constraints, any large-scalerestoration program should also consider prioritizing areasbased on restoring local ecosystem services (eg hydrologi-cal services and prevention of soil erosion) or protectingendemic biodiversity (eg through the creation of forestcorridors to connect isolated reserves), even if high levelsof active management are needed.

In an effort to account for such considerations, the

member organizations of the AFRP have developed mapsprioritizing areas for restoration based on the demands fordifferent ecosystem services, including the supply ofdrinking water to urban populations and suitable areas forcarbon sequestration projects. Opportunities for increas-ing landscape connectivity and low-productivity pastureson marginal sloping land to avoid competition for landwere also prioritized, as were potential areas for offsettingthe loss of native forests, according to the Brazilian ForestCode (Brancalion et al. 2013). Indeed, landholdings inthe Atlantic Forest must maintain 20% of native vegeta-tion cover as so-called Legal Reserve, and if this require-ment is not met, the deficit must be offset by restoringdegraded lands, or by acquiring or renting remnants ofequivalent size (Brancalion et al. 2013). Although moreecologically sensitive areas should be strictly protectedand should therefore be restored to meet conservationgoals, large-scale restoration elsewhere can benefit fromincreased revenues through sustainable management.Innovative models of forest restoration have been devel-oped to create incentives for farmers to invest in restoringdegraded areas, for example by allowing the sustainableextraction of both timber and non-timber forest products,coupled with payments for ecosystem services(Brancalion et al. 2012). The AFRP estimates that more

than three million local jobs could be generated as part ofthe restoration process (Brancalion et al. 2013), whileimprovement of degraded watersheds has the potential toreduce the cost of treating drinking water by a hundred-fold (Tundisi 2014).

Atlantic Forest and cattle pasture restoration in

Esprito Santo, Brazil

The Brazilian state of Esprito Santo provides a valuablecase study for understanding the potential challenges andsolutions involved in accommodating new restoration

areas in a region where land is an increasingly scarceresource. The state government, supported by both theState Institute for Environment and HydrologicalResources (IEMA or Instituto Estadual do MeioAmbiente e Recursos Hdricos) and the Agroforestry andForestry Defense Institute (IDAF or Instituto de DefesaAgropecuria e Floresta), has recently published the

Reforest Program (Reflorestar in Portuguese) withthe aim of restoring 236 000 ha of forest through large-scale restoration and conservation by the year 2025. Atthe same time, the state development plan calls for a284 000-ha expansion of the areas devoted to agriculturalcrops and a 400 000-ha expansion of forest plantations(PEDEAG 2008).

These proposed changes in land use take place within amosaic of existing agricultural landscapes, made up ofboth large and small landownerproducers. In 2010, thestate territory (4.6 million ha) consisted mainly of pas-turelands (41%, 1.9 million ha), croplands (19%, 862 000ha), native forests (21%, 990 000 ha), and silvopastoral

systems (11%, 487 000 ha; Lorena et al. 2014); other landuses include rocky areas (2.6%), other vegetation (2.3%),urban areas (1.5%), and water bodies (1.2%). Here, weassessed the potential carrying capacity of pastureland indifferent regions of Esprito Santo to accommodate684 000 ha of planned agricultural and forestry expansion(284 000 ha + 400 000 ha; Scenario 1) or the same agri-cultural and forestry expansion with an additional236 000 ha of native forest restoration (Scenario 2). Wethen examined how different increases in pasture produc-tivity could, in theory, provide new areas of land to satisfystate targets for both arable crops and restored forest.Pasturelands in the state were divided into polygons, andfrom these we extracted the potential forage grass biomassgrowth (kilograms per hectare), using a spatial databasedeveloped by the Food and Agriculture Organization andthe International Institute for Applied Systems Analysis(FAO/IIASA 2010). We converted biomass figures intoestimates of potential carrying capacity (AU per hectare)using standard coefficients for grazing efficiency (8 kgday1 dry biomass consumption per head and 50% pasturegrazing efficiency).

Assuming that beef production remained constant(PEDEAG 2008), we considered a target for improvingpasture productivity in a region to be feasible if two con-

ditions were met: (1) the increase in productivity is equalto or less than double the average productivity until theyear 2025, and (2) the final average stocking rate is lessthan or equal to 80% of the carrying capacity (the poten-tial number of animals per unit area; see WebPanel 1 fordetails). We viewed these as conservative, practical limitsfor changes in cattle productivity, given that we onlyconsidered fully grass-based systems; higher stocking ratescould be achieved with supplementary feed or confine-ment systems, but we did not consider these approachesbecause of animal welfare considerations. Our approachfor modeling land-use dynamics to account for the large-


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scale restoration of native ecosystems can easily adapted to other scenarios, including situations whebeef production is likely to increase.

We found that the potential carrying capacity of existinpastures in the state is 5.29 million AU, or 2.77 AU ha1. comparison, the current cattle herd in Esprito Santo is 1.4million AU (only 27% of the estimated capacity), with aaverage productivity of 0.74 AU ha1 (Figure 2 and Tab1). Most of the state is characterized by low levels of cattproductivity but with considerable potential for grow(Figure 3), further reinforcing the notion that increasi

cattle productivity in these areas is a viable option to spaother areas for restoration (Figures 4 and 5).

In Scenario 1, a 57% increase in cattle productivi(equivalent to an increase from 0.74 to 1.16 AU ha1)necessary to meet state goals for agriculture and foreplantations while maintaining current levels of beef prduction. In Scenario 2, a 93% increase in cattle produtivity would theoretically spare enough land to me

Figure 2. Map of the Brazilian Atlantic Forest (a) based on da

from SOS Mata Atlantica (2014). Areas in light green and dagreen show the original limit of the Atlantic Forest and its curreremnants, respectively. The state of Esprito Santo is marked w

a black outline. Current (b) and potential (c) pastuproductivity in Esprito Santo in AU ha1 are shown on a sc

where red represents low productivity (less than 0.5 AU ha

and blue represents high productivity (more than 3 AU ha1

).

(a)

(b) (c)


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agriculture and forestry expansion goals as well as torestore 236 000 ha of native forests over 15 years(between 2010 and 2025). This would mean reaching52% of the pasturelands carrying capacity in 15 years.

The risks of agriculture displacement (and thereforepotential for indirect land-use change and associatedsocial problems) following large-scale restoration vary

between regions within Esprito Santo. Competition forland between agriculture and forest restoration is likely tobe highest in the coastal northern region, Litoranea

Norte, which contains more high-productivity pasture-lands (Figure 2 and Table 1) and is home to a variety ofother highly profitable agricultural activities (Eucalyptus,coffee, and sugarcane). By contrast, competition for landis likely to be much less intense in the northwest region,given the areas relatively low levels of pastureland pro-ductivity (Figure 2 and Table 1).

Agricultural development, large-scale restoration,

and competition for land

Reconciling agricultural development, forest conserva-tion, and restoration is one of the greatest challenges forenvironmental and social sustainability in the face of cur-rent and future competition for land. If poorly managed,large-scale restoration like any externally motivatedand unfamiliar type of land use could result in unequaldistribution of benefits, leading to increased inequalityamong the original landowners and the potential dis-placement of more marginalized community members

(Barr and Sayer 2012). In addition, any loss of commodi-ties resulting from the original land use could be compen-sated by the expansion of new production areas else-where, if demand for those original commodities remains

high (ie indirect land-use change).Should this expansion occur in areascontaining old-growth vegetation,then this could result in negativeimpacts on biodiversity and ecosystemservices, even in situations wherethere is a net increase in forest cover(eg Ferraz et al. 2014).

The evidence for indirect land-usechange driven by the expansion ofland uses other than restorationremains ambiguous. While someauthors have demonstrated that soy-beans and sugarcane lead to displaceddeforestation in the Amazon (Arima

et al. 2011), others have been unableto show leakage of soybean expansioninto the Cerrado (a large area of sub-tropical and tropical savanna) of MatoGrosso after increased enforcement inthe Amazon (Macedo et al. 2012).Some studies found no evidence ofprotected area expansion in theAmazon causing deforestation else-where (Soares-Filho et al. 2010).There are also considerable method-ological challenges associated with

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Figure 3. Much of Esprito Santo is currently characterized bylow stocking rates (expressed in animal units per hectare on the xaxis) but high potential stocking rates (in animal units per hectareon the y axis), further reinforcing the major potential for increasing

cattle ranching productivity while sparing land for other uses.

Table 1. Summary of land-use indicators, based on the two scenariosanalyzed in this study

Scenario 2(increasing cattle

Scenario 1 ranching productivity (increasing cattle to increase croplandsranching productivity and forest plantationsto increase croplands and to meet

Land-use indicators Current and forest plantations) restoration targets)

Cropland area(thousand hectares) 862 1146 1146

Plantation forest area(thousand hectares) 487 887 887

Pastureland area(thousand hectares) 1909 1225 989

Stocking rate(AU per hectare) 0.75 1.16 1.44

Sustainable stockingcapacity (%) 27 42 52

Total native forest cover(thousand hectares) 990 990 1226

Restored forest area(thousand hectares) 0 0 236

Native forest coverincrease (%) 0 0 24

Potentialstockingrate(anim

alunitsperhectare)

Current stocking rate (animal units per hectare)


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assessments of leakage (Henders and Ostwald 2014).However, the potential for major re-allocations in landuse affecting vegetation clearance rates in neighboring

regions clearly remains a risk, especially in the context ofongoing transport and other infrastructure developmentsoccurring in the Amazon and Cerrado. For example, live-stock from the Atlantic Forest region has likely been dis-placed to the Amazon and Cerrado after the expansion ofmore profitable land uses such as sugarcane (Lapola et al.2014), and absolute deforestation rates in the Cerradorecently surpassed those in the Amazon, where environ-mental enforcement is more rigorous (Soares-Filho et al.2014). As such, restoration efforts in the Atlantic Forestat the scale of millions of hectares could possibly have asimilar effect, resulting in further clearance of old-growthfragments in the Atlantic Forest or in the neighboringCerrado.

Sustainable increases in production on current agricul-tural lands are commonly suggested as a key part of effortsto ease the conflict between agricultural expansion andthe conservation of natural ecosystems (Smith et al. 2010;Latawiec et al. 2014a); increases in cattle ranching pro-

ductivity often provide the most promising opportunito spare large areas of land from deforestation (eg Cohnal. 2014; Strassburg et al. 2014b). Nevertheless, the imp

cations of a land-sparing approach with respect to largscale restoration have been poorly explored. Here wshow that in the case of the Atlantic Forest, plausibincreases in cattle productivity, the dominant and leaproductive land-use type in the region, could free uenough land to meet large-scale forest restoration targein Esprito Santo, helping to restore critical ecosysteservices while providing positive returns for landownerincluding small-scale cattle ranchers and the widdevelopment of the regional economy.

Yet there are several essential pre-conditions for thsuccess of any such land sparing, with coupled improvments in agricultural productivity (whether in cattranching, as in this case, or another large-scale agricutural system) and restoration. First, restoration efforts animproved agricultural practices incur start-up costs anrequire necessary technical support and specializknowledge, especially in areas that are highly degraded are isolated from native forest. Even though these inve

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Figure 4. Restoration planting in an area formerly used for cattleranching in Rio de Janeiro state, in the Atlantic Forest region. (a)

The area 3 months after the introduction of native tree seedlingsto degraded pastureland; (b) the same area 1 year later.

Figure 5.An increase in pasture productivity in areas suitable for cattle ranching (left) allowed a farmer to set aside marginal arewith rocky soils (right) for forest restoration in the Atlantic Forest in Itu-Sao Paulo, southeastern Brazil.

(b)(a)


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ments may be recovered relatively quickly (eg 34 years;Brancalion et al. 2012), there are often problems with ini-tial financing as well as with the educational and culturalbarriers involved in adopting more technologicallyadvanced agricultural systems (Rodrigues et al. 2011;Brancalion et al. 2012; Latawiec et al. 2014b).

Second, improvements in the benefits and profitability

of both agriculture and restoration forestry need to beclosely monitored and integrated across entire propertiesand landscapes, especially in areas where competition forland is strong. Restoration forestry needs to become aneconomically viable land use, where the exploitation oftimber and non-timber forest products, as well as the pos-sibility of payments for ecosystem services from restora-tion sites, could provide returns that are comparable tothose from the preceding agricultural activities (egBrancalion et al. 2012; WebPanel 2).

Third, any intervention to improve the profitability ofone major land use (eg cattle ranching) must be accompa-nied by effective regulatory policies enforcing strict pro-

tection for remaining areas of native vegetation. Withoutsuch measures, increased productivity can easily lead toincreased deforestation (ie the rebound effect, whereincreased productivity and hence profitability leads toan increase in demand for more land and still greater pro-ductivity; Meyfroidt and Lambin 2009; Rudel et al. 2009).Improvements in the profitability of agriculture may alsoescalate future costs of conservation and restoration activ-ities as increases in land rents outpace investments in con-servation incentives (Phelps et al. 2013).

Finally, improvements in the productivity of one agricul-tural sector must not result in negative social consequences:for example, disenfranchising smallholders followingincreases in land prices, failing to incorporate originallandowners in any process of technological improvement,or neglecting to protect the livelihoods of those involved inother, less-profitable farming activities (eg staple crop pro-duction, fruticulture, and agroforestry) that are key to meet-ing local and regional food security needs.

An integrated suite of policies should guarantee thatlarge-scale restoration delivers long-term environmentaland social benefits: strategic territorial planning (egthrough Brazils economicecological zoning plans),improved enforcement of existing environmental regula-tions, land-tenure security, monitoring of land-use prac-

tices, incentives for job provision through restorationwork, and other social welfare and justice considerations(Calle et al. 2012). Strategic planning for enhanced land-scape connectivity and prioritizing restoration in areas ofhigh conservation value must also be taken into accountif large-scale restoration is to deliver long-term benefitsfor biodiversity (Banks-Leite et al. 2014). Although wehave focused here on cattle production, the risks andopportunities associated with large-scale restoration arerelevant to other agricultural systems as well. There maybe situations, even in areas dominated by extensive cattleranching, where displacement effects can be avoided by

switching to another production system. Indeed, naturalresource managers should consider the range of activitiesand approaches that may contribute toward the improveduse of any system, including diversification and process-ing of commodities. It then becomes possible to set asideareas for large-scale restoration, thus ensuring the protec-tion of both local and regional ecosystem services while

avoiding potential negative displacement.

Acknowledgements

AEL and BBNS acknowledge Conservation International,the Gordon and Betty Moore Foundation, and the

Norwegian Agency for Development Cooperation(Norad) for funding. TG acknowledges the SwedishResearch Council Formas (Grant 2013-1571) for funding.We thank M Simas and A Iribarrem for help withmanuscript preparation, M Rangel for the photograph inFigure 1, F Barros for help with the map of Atlantic Forestand Figure 3, and F Cronemberger for help with panels

b and c in Figure 2. FR Scarano, LP Pinto, and M Sossai arealso acknowledged for their help throughout the durationof the study.

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