REVIEW SUMMARY

CONSERVATION

Landscapes that work for biodiversityand peopleC Kremen and A M Merenlender

BACKGROUND Biodiversity is under siegewith greatly enhanced rates of local and globalextinction and the decline of once-abundantspecies Current rates of human-induced climatechange and landuse forecast theAnthropoceneas one of the most devastating epochs for lifeon earth How dowe handle the Anthropocenersquostriple challenge of preventing biodiversity lossmitigating and adapting to climate change andsustainably providing resources for a growinghuman population The answer is in how wemanageEarthrsquos ldquoworking landsrdquo that is farmsforests and rangelands These lands must bemanaged both to complement the biodiversityconservation goals of protected areas and tomaintain the diverse communities of orga-nisms from microbes to mammals that con-tribute to producing food materials cleanwater and healthy soils sequestering green-house gases and buffering extreme weatherevents functions that are essential for all lifeon Earth

ADVANCES Protected areas are the corner-stone of biodiversity conservation Althoughthe total area of protected regions needs to beincreased parks will nonetheless continue tolose species if these areas are isolated fromoneanother by inhospitable land uses and arefaced with a rapidly changing climate Furthermany species such as those that migrate re-main unprotected as they occupy lands outside

of parks for all or portions of their life cyclesLastly protected-area effectiveness is greatlyinfluenced by surrounding landmanagementldquoWorking lands conservationrdquo aims to sup-port biodiversity while providing goods andservices for humanity over the long term assur-ing sustainability and resilience By manag-ing lands surrounding parks favorably workinglands can buffer protected areas from threatsand connect them to one another This ap-proach complements protected areas by pro-viding accessory habitats and resources forsome species while facilitating dispersal andclimate change adaptation for others Furtherby maintaining the biodiversity that suppliescritical ecosystem services within workinglands these approaches ensure that the pro-duction of food fiber fuel and timber can besustained over the long run and be more resil-ient to extreme events such as floods droughtshurricanes andpest anddisease outbreaks whichare becomingmore frequent with climate changeA variety of biodiversity-based land managementtechniques can be used in working lands includ-ing agroforestry silvopasture diversified farmingand ecosystem-based forest management to en-sure sustainable production of food and fiber

OUTLOOK The underlying principle ofbiodiversity-based management of workinglands has been practiced since ancient timesToday these systemshave largely been replaced

by unsustainable resource extraction ratherthan serving as models that could be adaptedto modern conditions Although various reg-ulatory voluntary and financial tools exist topromote sustainable landmanagementmanybarriers prevent individuals communities andcorporations fromadopting biodiversity-basedpractices including deeply entrenched policyand market conditions that favor industrializedor extractive models of land use Thus uptake

of these approaches hasbeen patchy and slow andis not yet sufficient to cre-ate change at the tempo-ral andspatial scalesneededto face the triple Anthro-pocene threat

Biodiversity-based land management prac-tices are knowledge- rather than technology-intensive They are well adapted to empowerlocal communities to manage their naturalresources One of the most exciting emergingtrends is community-driven initiatives toman-age working landscapes for conservation andsustainability By linking up through grass-roots organizations social movements andpublic-private partnerships these initiativescan scale up to create collective impact and candemand changes in government policies tofacilitate the conservation of working landsScientists and conservation practitioners cansupport these initiatives by engaging with thepublic listening to alternativeways of knowingand cocreating landscapes that work for bio-diversity and people

RESEARCH

Kremen et al Science 362 304 (2018) 19 October 2018 1 of 1

TOMORROWrsquoS EARTHRead more articles onlineat scimagTomorrowsEarth

The list of author affiliations is available in the full article onlineCorresponding author Email ckremenberkeleyeduCite this article as C Kremen and A M Merenlender Science362 eaau6020 (2018) DOI 101126scienceaau6020

Strawberry production inCentral Coast CaliforniaOnthe left a homogeneous land-scape of strawberry mono-culture including organic fieldssupports fewer wild speciesthen a diversified organic farm(right) in the same region whichincludes a small field of straw-berry surrounded by orchardshedgerows diverse vegetablecrops and natural habitatsThemonoculture landscape createsbarriers to wildlife dispersalwhereas the diversified land-scape is more permeable

PHOTOCKREMEN

ON OUR WEBSITE

Read the full articleat httpdxdoiorg101126scienceaau6020

on October 18 2018

httpsciencesciencem

agorgD

ownloaded from

REVIEW

CONSERVATION

Landscapes that work for biodiversityand peopleC Kremen and A M Merenlender

How can we manage farmlands forests and rangelands to respond to the triple challengeof the Anthropocenemdashbiodiversity loss climate change and unsustainable land useWhen managed by using biodiversity-based techniques such as agroforestry silvopasturediversified farming and ecosystem-based forest management these socioeconomic systemscan help maintain biodiversity and provide habitat connectivity thereby complementingprotected areas and providing greater resilience to climate change Simultaneously theuse of these management techniques can improve yields and profitability more sustainablyenhancing livelihoods and food security This approach to ldquoworking lands conservationrdquo cancreate landscapes that work for nature and people However many socioeconomic challengesimpede the uptake of biodiversity-based land management practices Although improvingvoluntary incentives market instruments environmental regulations and governance isessential to support working lands conservation it is community action social movementsand broad coalitions among citizens businesses nonprofits and government agencies thathave the power to transform how we manage land and protect the environment

Biodiversity the product of 38 billion yearsof evolution is under siege Not only areboth marine and terrestrial species expe-riencing accelerated rates of local andglobalextinction (1ndash3) but even common species

are declining (2 4 5) This alarming situationhas prompted a strong call for increasing thenumber (6 7) and effectiveness (8) of protectedareas the principal method for combatting spe-cies loss Though suchprotections are essential wecannot rely on protected areas alone to preservespecies As protected areas become increasinglyisolated because of habitat loss and degradationmuch research has revealed that they will lose spe-cies over time (9) Furthermany critical threats tospecies do not respect protected-area boundaries(10) including climate change which both exac-erbates species losses (11) and threatens to alterthe biomes of many currently protected regionsentirely (12)More hopefully recent studies show that some

human-dominated landscapes can supportmuchmore biodiversity than previously recognized(13ndash17) suggesting a complementary path for-ward Specifically when these areas generallyreferred to as the ldquomatrixrdquo represent a high-quality mosaic of land uses they can play a crit-ical role in sustaining biodiversity both in situand by promoting species dispersal among pro-tected areas and remnant habitats and alongmigratory routes (Fig 1) (15 18 19) Of coursehuman survival also depends on the long-termcapacity of this matrix of ldquoworking landsrdquo in-

cluding rangelands forests and farms to pro-duce food water fiber fuel and forest productsAll too often however these goods are producedat severe environmental cost including habitatdegradation toxic contamination and depletionof water quantity and quality leading to ecologi-cal collapse local extinctions and the creation ofunproductive wastelands (20 21) We argue thatinstead working lands can be used to supporthigh levels of biodiversity while satisfying humanneeds in a sustainable way Because rangelandsforests and cultivated lands collectively occupy~80 of terrestrial area (21) the potential for con-servation in such lands is enormousCritical ecosystem functions and services are

provided by a suite of diverse organisms frommicrobes to mammals and thus maintenanceof these organisms is necessary for long-termand sustainable productivity of working lands(22 23) Hence managing the matrix to main-tain biodiversity is not only necessary for speciesconservation but also essential for sustainable pro-duction Biodiversity-based production systemsincluding agroecological farming or ecosystem-based forest management are often perceived as

unproductive an incorrect viewpoint that im-pedes the public investment needed to developand promote these methods Here we describemanaging the matrix jointly and sustainably forbiodiversity and people through ldquoworking landsconservationrdquo and askwhat strategies can be usedto strengthen and scale up this approach asrapidly as possible to help combat the tripleAnthropocene threats of biodiversity loss cli-mate change and unsustainable land use

Working lands conservation defined

Although the term ldquoworking lands conservationrdquois already used in policy statements and in guid-ance for conservation programs [eg (24)] theconcept has yet to be formally defined and risksbeing misapplied We define it at the landscapescale (Box 1)To avoid mass extinction and ecosystem col-

lapse we must integrate biodiversity conserva-tion into the landscapes we use and not simplyrelegate nature to a limited number of protectedareas that are doomed if left as isolated habitatislands within biological deserts Working landscan provide food breeding sites and shelter fora myriad of species while maintaining abioticconditions including temperature light windwater fire and other disturbance processeswithin required ranges They can facilitate func-tional connectivitymdashthat is themovement of orga-nisms across the landscape and among habitatpatches that promotes population persistence byallowing for gene flow recolonization and adap-tation to climate and other global changes (25 26)To support humanity sustainably a working

landscape must be productive and maintain theecosystem services such as pollination pestcontrol and nutrient cycling that underlie thatproduction Maintaining these services requiressupporting the underlying populations of service-providingorganismsWithin each service a greaterdiversity of service providers often enhances thelevel andor quality of services and reduces un-certainty in service delivery (22) because differentspecies respond differentially to environmentalchange (27 28) Maintaining connectivity is alsoimportant both to support flows of ecosystemservice providers andormaterials (eg pollinationrequires animal vectors to move pollen betweenflowers water purification requires water to flowthrough vegetation) (29) and to enhance meta-community persistence of service-providing orga-nisms to sustain ecosystem functions and servicesover space and time (22 30)

RESEARCH

Kremen et al Science 362 eaau6020 (2018) 19 October 2018 1 of 9

Department of Environmental Science Policy andManagement University of California Berkeley BerkeleyCA 94720 USACorresponding author Email ckremenberkeleyedu

Box 1 Definition of working lands conservation

DefinitionConservation inworking landscapesmaintains biodiversity provides goods and servicesfor humanity and supports the abiotic conditions necessary for sustainability and resilience

These socioecological systems both support biodiversity by providing critical resources andrely on biodiversity (specifically ecosystem service providers) for sustainable production offood water fiber fuel and forest products These landscapes also enhance connectivity topromote the movement of organisms natural processes and ecosystem services

Working lands conservation emphasizes the critical role of managing the matrix for speciesconservation to complement protected areas

on October 18 2018

httpsciencesciencem

agorgD

ownloaded from

Ensuring the sustainability of production re-quires balancing across provisioning regulat-ing and supporting services in other wordsseeking multifunctionality and stability ratherthanmaximal production For example conven-tional (chemically intensive) monoculture agri-culture produces high yields but often at theexpense ofwater quality climate regulation andsoil health (Fig 2A) (20) and can suffer produc-tion collapse in response to periodic extremeweather pests and diseases (31ndash33) Althoughtransforming to a more sustainable systemmayreduce average yields somewhat [eg (34)] byrelying on ecosystem services produced on thefarm and in the surrounding landscape a sus-tainable system is bothmultifunctional andmoreresilient to change (20 31) (Fig 2C)Working landscapes often comprise hetero-

geneous patch types including novel commu-nitiesmade up ofmixtures of native andnonnativespecies as well as remnants of natural or semi-natural habitats whose composition is more simi-lar to that of a historical ecological community(35) Althoughmanagement goals likely will dif-fer among patch types both individual patchesand the whole landscape should be managedfor sustainability For example patches whosecommunities are far fromhistorical could beman-aged principally for crops (a provisioning service)by using sustainable agricultural practices tomin-imize negative effects on biodiversity and ecosys-tem services on and off site Remnant patchescould be retained as stepping-stone habitats to

support species dispersal and provide regulatingservices such as pollination (29 31) Maintainingmosaic landscapes composed of different patchtypes provides opportunities to maximize diver-sity resilience andmultifunctionality Radar dia-grams reveal likely trade-offs and sustainabilitywithin and across patches (Fig 2B) as well asmultifunctionality at the landscape scale (Fig 2C)Conservation in working landscapes draws

upon several related concepts Integrated land-scape management initiatives seek to simulta-neously improve food production biodiversityor ecosystem conservation and rural livelihoodsand are being implemented by governments andnongovernmental organizations in Latin Americaand Africa (36) The ecosystem stewardship con-cept focuses on the need to sustain Earthrsquos capac-ity to provide ecosystem services and supportsocioecological resilience under conditions ofuncertainty and change (27) The socioecologicalproduction landscape of the Japan SatoyamaSatoumi Assessment refers to dynamic landscapemosaics that have been shaped over time by theinteractions between people and nature in waysthat jointly support biodiversity and humanwell-being (37) These concepts also emphasize criticalsocial components such as involving multiplestakeholders at the landscape scale communityparticipation intersectoral coordination flexibleand adaptive governance systems social learn-ing and adaptive management which are nec-essary for successful conservation of workinglandscapes

The underlying principle of maintaining eco-logical diversity inherent to these approacheshas been practiced since ancient times Some ofthese management systems such as indigenoususe of fire weeding pruning and the seed dis-persal that shapedCalifornian ecosystems (38) nolonger exist in their original formwhereas otherssuch as regional pastoral andhigh-mountain farm-ing systems in Europe (39) persist in some areasBy creating highly simplified and intensified pro-duction systems (21 40) from corn and soy inUSmidwestern states to palm oil plantations insoutheast Asia and vineyards in Chile we haveabandoned this critical sustainability principleacross much of Earthrsquos cultivated landscapesHowever it is a fallacy that such systems willultimately spare more land for nature conser-vation or feed the world indefinitely rather weneed to find ways to allow biodiversity-basedproductionmethods to figuremuchmore prom-inently in local regional and globalmarkets (16)

Working lands conservation as acomplement to protected areas

Given the dire situation facingmany species andthe expectation of further species losses and shiftsin ecosystem composition due to climate change(2 4 11) ceasing further habitat conversioncompletely and protecting large regions of Eartheffectively are critical necessities for conservation(6ndash8) although just how much should be pro-tected is highly debated (41) [By ldquoprotected areardquowe refer to parks whose primary function is toconserve biodiversity and wilderness (Interna-tional Union for Conservation of Nature andNatural Resources categories I to IV constituting675 of terrestrial area) (42) in contrast to areasblending conservation and livelihood objectives(categories V to VI constituting 865)] How-ever the protected-area strategy alone will notbe successful without complementary workinglands conservation in the surrounding landscapesFirst even the largest protected areas will losespecies over the long term (9) unless surroundinglandscapes can be managed to provide connec-tivity among parks Further less than 10 of pro-tected areas are expected to represent currentclimatic conditions within 100 years increasingthe criticality of matrix connectivity to permitspecies to follow their suitable climates (12)Lastly effectiveness in controlling threats suchas invasive species encroachment poaching andother impacts on protected lands also criticallydepends on the surroundingmatrix (10) Thus tostem the tide of biodiversity loss wemust expandbeyond protected areas using working lands con-servation both to buffer and to reduce the threatsthat cross park boundaries and to create acces-sory habitats for bothmovement and persistenceWorking lands conservation is a key linchpin

for combatting the triple Anthropocene chal-lenge of biodiversity loss climate change andunsustainable land use A large-scale example isthe Mesoamerican Biological Corridor projectwhich has fostered a multistakeholder partic-ipatory process to enhance connectivity on culti-vated range and forest working lands to link

Kremen et al Science 362 eaau6020 (2018) 19 October 2018 2 of 9

Fig 1 Rebuilding connectivity in the matrix by using silvopasture Photo of Finca La Luisashowing several types of silvopastoral systems including regenerating secondary tropical dry foresttrees with grass understory (yellow) and rows of planted Eucalyptus trees interspersed withnitrogen-fixing Leucaena leucocephala fodder shrubs and forage grasses (blue) These systems wereestablished on former monoculture agricultural lands to restore compacted degraded soils thered area shows early stages of tropical dry forest regeneration prior to grass seeding for silvopastureSilvopastures produce more cattle sustainably on less land buffer ranchers from economic lossesdue to climate extremes and create landscape connectivity to other forest fragments (orange) in theCesar river valley Colombia

RESEARCH | REVIEWPHOTOJ

JLO

PERAC

IPAVADAPTED

BYNIRJA

DESAISCIENCE

on October 18 2018

httpsciencesciencem

agorgD

ownloaded from

more than 650protected areas in the region (43)A concurrent goal is to use sustainable agricultureand forestry techniques to promote livelihoodsand enhance resilience to climate change (36)Protected areas are vital in this region becausemany species are restricted to forest howevermostreserves are small and isolated In combinationwith steep elevational and latitudinal gradientsin the region this isolation makes species in-habiting reserves particularly vulnerable to climatechange The Mesoamerican Biological Corridorproject recognizes the role thatworking lands canplay to restore critical connectivity by increasingtree diversity and cover through live fences agro-forestry silvopasture forest fallows homegardensand protection or restoration of riparian forestsand forest fragments (43) These forest elementswhich include both ribbonlike and patch struc-tures support a large number of neotropicalbirds insects mammals and plants (17 44) en-hance themovement of birds and bats across thelandscape (45ndash47) and thus contribute to con-servation even of vulnerable wildlife (17 47 48)Forest elements also promote sustainable landuse and contribute to local livelihoods by sup-

porting ecosystem services For example evi-dence suggests that an economically devastatinginvasive pest the coffee berry borer is reduced bythe integration of forest elements within coffeelandscapes which both limits the borerrsquos abilityto colonize new coffee fields (49) and promotesbird species that prey on the borer (50) Reducedeconomic losses due to pest control frombirds aresimilar inmagnitude to average per capita incomein the region and are strongly related to forestcover (50) Adopting sustainable agriculturaltechniques and enhancing tree cover simulta-neously creates more flexible and resilient pro-duction systems that allow farmers and ranchersto adapt to extreme conditions prompted byclimate change (33 51) Although some criticsdecry the effectiveness of the MesoamericanBiological Corridor project itmay be too early tojudge Quite a few integrated landscape initiativesare concentrated in the region in associationwithbiological corridors (36) However many beganrelatively recently and we know from the fewscientific studies that exist that developing aneffective multistakeholder participatory processtakes substantial time (36 43 52) In one case

that is more advanced (the San JuanndashLa SelvaBiological Corridor in Costa Rica) some successhas been achieved in arresting deforestation andencouraging tree planting forest regenerationand connectivity through a government-run pay-ments for ecosystem services program as well asother grassroots initiatives (43 53)

Mechanisms for promoting workinglands conservation

The challenge of shifting frommanaging work-ing lands solely for profit to conservation ofworking lands is not insignificant but there areclear paths toward larger-scale integration ofthis approach These strategies include variousregulatory voluntary incentive market-basedor governance instruments (table S1) which varyin their applicability to private communal orstate-owned lands and the extent to which theysupport biodiversity conservation versus liveli-hoods or economies (Fig 3A) Each approach haschallenges especially around reconciling conser-vation and socioeconomic objectives (table S1)(42 54) Collectively problems associated withregulatory and incentive programs can includeinter alia lack of permanence or compliance com-plex implementation unintended economic con-sequences low adoption rates high monitoringcosts and little evaluation of effectiveness againstgoals (table S1)Further there is often the risk that the bio-

diversity conserved through these actions is notequivalent to that which was lost because of eco-nomically driven land conversion Instrumentsfor private lands may result in piecemeal landmanagement actions that have little positive ef-fect on biodiversity at the landscape scale promis-ing public-private initiatives to overcome thisdefect include corridor planning (43 55) (Box 2and Fig 4) and landscape-levelmitigation (tableS1) For example landowners required to set asideforest on their properties under Brazilrsquos forestcodemay develop these lands in exchange formit-igating lands elsewhere within the same biomethat provide greater conservation value (56)Man-aging the matrix to promote biodiversity couldalso exacerbate human-wildlife conflict how-ever the recovery of carnivore populationswithinhuman-dominated areas in Europe provides ahopeful and inspiring example for how landscapescan be shared between wildlife and people (14)(Box 3) These instruments can exacerbate theunequal distribution of benefits and costs withinand across communities (table S1) For exampletrading development rights on forestlands in ex-change for permitting high-density urban devel-opment elsewhere can provide open spaces forworking lands conservationHowever such tradescould exacerbate the lack of access to open spacealready experienced by low-income urban house-holds Thus the effects of conservationmeasureson social equity and environmental justice shouldalso be considered (57) A final concern is thatthere is often a trade-off between the rigor ofenvironmental standards or restrictions enforcedand the likelihood of adoption (table S1) incen-tive schemes that are flexible provide obvious

Kremen et al Science 362 eaau6020 (2018) 19 October 2018 3 of 9

A

B

C

Crops

Healthysoils

FreshwaterCarbon sequestration

Pest control services

Pollination services

Biodiversity

Connectivity

Forest products Livestock

production

Biodiversity Crops

Healthysoils

Freshwater

Carbon sequestration

Pest control services

Pollination services

Connectivity

Forest products Livestock

production

Monoculture row-crop Mixed cultivated forest and range landscape

Rangelands

Diversified farm

Riparian forest

Fig 2 Ecosystem service trade-offs with land management Radar diagrams display howdifferent land uses affect various ecosystem services and biodiversity (A) Monoculture row croppingcontributes to food production at the expense of other ecosystem services and biodiversity (B) In aworking landscape managed for conservation patch types differ in the services they provide but eachpatch type should display a relatively even array of services minimizing trade-offs (C) Acrosspatches the services provided for the working landscape in (B) are multifunctional

RESEARCH | REVIEWILLU

STRATIO

NNIRJA

DESAISCIENCEBASED

ON

CKREMEN

AND

AM

MERENLE

NDERPHOTONATIO

NALAERIALIM

AGERYPROGRAMADAPTED

BYNIRJA

DESAISCIENCE

on October 18 2018

httpsciencesciencem

agorgD

ownloaded from

benefits target likely adopters fit the sociocul-tural context foster enabling market and reg-ulatory environments and provide technicalassistancemay boost adoption (58) For examplepayments for conserving or restoring forests inCostaRica are based on area whereas transactioncosts are the same regardless of size disincen-tivizing smaller landowners from participatingin the payments for ecosystem services schemeEncouraging smallholders to participate wouldrequire adjusting the costs of participation sothat these landowners could also realize net gains(53) Although numerous changes are requiredcareful attention to the construction of theseprograms could increase their successFurther several current trends favor working

lands conservation approaches First new policyinstruments [such as REDD+ (Reducing Emis-sions from Deforestation and Forest Degrada-tion)] operating across a range of scales fromindividual private landholdings to large-scalecommunity-based or government-funded initia-tives are being developed to incentivize conser-vation onworking lands Second the number andvariety of institutions involved in working landsconservation are increasing and such institu-tions include both public-private partnershipsand nongovernmental conservation organiza-tions that formerly focused primarily on pro-tected areas (36 59 60) Third these institutionscan take advantage of recent increases in bothpublic and private ldquoinvestments for conserva-tionrdquo (investments designed to cogenerate finan-cial returns and conservation benefits) (60) Suchinvestments include projects in sustainable foodand fiber production water quality and quantityprojects and outright habitat conservation (inthe latter financial returns are based on chang-ing land values or carbon stocks) Fourth out-side of these investments an increasing numberof companies have committed to greening theirsupply chains by reducing the environmentalimpacts at the source processing delivery andend-of-life management of the product (61) Al-though supply chain greening requires muchbetter monitoring accountability and inclusionof biodiversity conservation as an explicit goal(61 62) it could ultimately contribute to conser-vation in working landscapes particularly giventhe vast economic power represented within cor-porations (61) A final trend is the creation ofvoluntary community-driven programs (Box 2)in which local communities participate in theconservation of working landscapes to gain in-creased access to information and expertise buildinterpersonal connections and obtain both per-sonal benefits and public recognition for practic-ing sustainable methods (63)We argue that this latter trend of community-

based actions and the innovations networksand social movements that sometimes emergefrom thempresent themost exciting opportunityto turn the tide against the triple Anthropocenethreat [see also (64)] Communities seeking solu-tions for socioecological resilience frequently relyon working lands conservation approaches Forexample Sustainable Solutions restores man-

grove forests in Sri Lanka and India throughyouth-based community engagement to buildshoreline resilience to cycloneswhile enhancinglivelihoods from fisheries dependent on man-grove ecosystems

Further local initiatives can link together toform larger networks with the help of boundaryorganizations to form socialmovements that canadvance environmental policies improve sus-tainable behaviors and demand supply chain

Kremen et al Science 362 eaau6020 (2018) 19 October 2018 4 of 9

Private

Public

Intensive agricultureConservation easement Private

reserveManagement incentives

Certification schemes

Mitigation

Payment for ecosystem services

Community based natural resources management

Indigenous conserved areas

Protected areas (IUCN Cat VVI)

Government landsGovernment plantation

Protected areas (IUCN Cat IndashIV)

Development

Cultivated landsLocal diversification and landscape-scale heterogeneity

Single-species patures

Rangelands (lt10 tree cover)

Forest lands

Conservation

A

B

Multi-species pastures

Row crop monoculture

Simple rotation

Monoculture tree plantations(even-aged) (multi-aged)

Silvo-pastoral system

Mixed croplivestock

Complex rotationintercrop

Management intensive grazing

Restored range

Nomadic pastoralism

Agroforestry system

Home garden

Nativenon-native multi-age forests

Restored forest

Native forest with EBM

Chemical intensification

Fig 3 Approaches for conservation of working lands occupy the space (yellow) between highlydeveloped (brown) and highly conserved (green) land uses (A) An array of tools are availablefor working lands conservation for private communal or public lands (see table S2 for more detail andexamples) IUCN Cat International Union for Conservation of Nature and Natural Resources categories(B) Forms of management for forage crops and tree products from cultivated lands (yellow)rangelands (light green) and forests (dark green) arrayed roughly along a management gradient ofdiversification (left to right) or chemical intensification (right to left) Cultivated lands include allplanted systems Dashed lines indicate overlapping concepts EBM ecosystem-based management

Box 2 Community stewardship The case of Landcare Australia

The Landcare movement is a well-documented community stewardship effort begun in themid-1980s to conserve biodiversity and sustain agriculture in Australia resulting in more than5000 Landcare and Coastcare groups More than 20 countries have since adopted the modelIn Australia this model combines substantial government investment with landowner andcommunity engagement For example Landcare groups across eastern Australia contribute tothe delivery of the Great Eastern Ranges (GER) Initiative (105) alongside public land managementauthorities conservation organizations research institutions and traditional owners groupsThe GER is one of Australiarsquos largest public-private partnerships to conserve biodiversity in theface of climate change (Fig 4) as part of Australiarsquos National Wildlife Corridors Plan Landcaregroups along the corridor undertake restoration and management activities along with communitybuilding and engagement In the Queanbeyan Landcare group 25 landholders signed up toincrease the foraging habitat for the glossy black cockatoo (Calyptorhynchus lathami) throughthe restoration of 10000 she-oaks (Allocasuarina sp) in production lands along three rivercatchmentsThe social networks and learning spaces created are promising ways of encouragingconservation commitment among land managers However far more landowners must becomeengaged to restore connectivity at the scale desired

RESEARCH | REVIEWILLU

STRATIO

NNIRJA

DESAISCIENCEBASED

ON

CKREMEN

AND

AM

MERENLENDER

on October 18 2018

httpsciencesciencem

agorgD

ownloaded from

accountability (64) For example the withdrawalof the United States from the Paris Agreementat the 21st Conference of Parties (COP21) anddelays in regulation of emissions by other nationsgalvanized a series of on-the-ground climateactions from civil society businesses nonpro-fits and subnational government The GlobalAction Climate Summit of 2018 instigated byCalifornia governor Jerry Brown illustrates anew stage of this growing social movement

Its Land and Ocean Stewardship ldquo30 times 30rdquo chal-lenge brings together more than 100 organ-izations focused onmanaging forests farmlandsand oceans to provide 30 of the climate changesolution by 2030 rather than waiting on agree-ments among nation states that continue tofall short of the necessary carbon reductiontargets The land management techniquesbeing developed locally to mitigate and adaptto climate change are generally consistent with

the conservation of working lands approach[eg (65)]The benefits of local land conservation can

also be scaled up andmademore effective if theyare carried out within a landscape or regionalconservation program organized by a state ornonprofit agency (58) Innovative social andinstitutional arrangements for working landsconservation may emerge such as The NatureConservancyrsquos BirdReturns program inCaliforniaThrough a reverse auction the program finds andpays farmers willing to alter water managementto create ldquopop-uprdquowetlands to provide habitatsfor shorebirds during their northward migra-tion selecting sites that optimize the conserva-tion benefits relative to payments (15)

Management techniques for conservingworking landsCultivated lands

Cultivated lands make up 12 of the terrestrialice-free surface (66) and comprise row and foragecrops seeded pastures vineyards and orchardsmixed crop and livestock systems and tree cropsand plantations (Fig 3B) Cultivated lands areoften highly simplified ecologically thus theyrely extensively on chemical fertilizers and pes-ticides to replace ecosystem services formerlygenerated within or around agroecosystems (31)often creating negative consequences for theenvironment and human health (Fig 2A) (21)including continued large-scale forest conver-sion in some areas of the biodiverse tropics (62)Instead diversified farming systems using ag-roecological management practices operate byfostering biophysical conditions and ecolog-ical interactions favorable to crop production(31 67 68) producing amore balanced (sustain-able) distribution of ecosystem services (Fig 2B)Evidence also suggests that they minimize manyof the negative environmental consequences as-sociated with simplified farming (31) (Fig 5) Fur-ther these techniques can maintain crop yieldsandprofitability create newmarket opportunitiesenhance food security nutrition and livelihoodsand contribute substantially to the global foodsupply particularly under a changing climate(table S2) Because they rely on relatively low-cost low-technology knowledge-basedmethods(69) agroecological diversification techniques canbe made accessible to the majority of farmers[Small-scale farms with lt5 ha make up 94 offarms worldwide (40) and produce more thanhalf of world food crops (70)] These farmingmethods use open-pollinated seed varieties thatcan be saved and cultivars that are locally adaptedthus they are less dependent on purchased seedsand other inputs that can lead to poverty traps(71) Multiple grassroots organizations and socialmovements support learning sharing and adapta-tion of agroecological knowledge and seedsthrough farmer-to-farmer networks under par-ticipatory governance (64) Diversified agroeco-logical practices are therefore farming methodsthat are highly compatible with working landsconservation although potentially more ap-plicable to certain farming systems Large-scale

Kremen et al Science 362 eaau6020 (2018) 19 October 2018 5 of 9

The great eastern ranges corridor Hinterland bush links

Australia

Border ranges alliance

Jaliigirr biodiversity alliance

Hunter valley partnership

Illawarra to shoalhaven

Southern highlands link

Kosciuszko to coast

Kanangra boyo to wyangala link

Slopes to summit

Central victoria biolinks alliance

1

2

3

4

5

6

7

8

9

10

Fig 4The GER Corridor Initiative AustraliaThe light green outline represents the plan to protectand restore more than 3600 km2 as a climate corridor The numbered dark green shapes denoteregional alliances of conservation and natural resource management organizations includingLandcare communities (Box 2) In the photo members of the Molonglo Catchment Group Landcarecommunity conduct restoration

Box 3 Carnivore conservation in shared landscapes

Maintaining populations of large carnivores ranks among the greatest of conservationchallenges These area-demanding species require larger territories than most protected areaspossess potentially necessitating costly translocations to ensure gene flow and maintainpopulations Further these species conflict with people in surrounding matrices throughpredation on livestock or occasionally maiming or killing of humans Nonetheless in Europemost large carnivore populations are stable or expanding One-third of the area of mainlandEurope hosts at least one permanent population of its four large carnivore species persistingalongside moderate human densities and largely outside of protected areas The success ofcarnivore conservation in Europe is attributed to well-enforced coordinated legislative protectionimprovements in habitat and ungulate prey base and rural depopulation Importantly ranchershave found ways to live with carnivores by using carnivore-proofed electric fences and re-invigorating traditional livestock-guarding practices using shepherds and dogs (14) Similarly in acultivated region in India large carnivore species (the leopard and striped hyena) persist withfew conflicts despite high human densities (300 peoplekm2) and the lack of wild prey (106)suggesting the potential that exists for carnivore conservation in shared landscapes

RESEARCH | REVIEWMAPSGREATEASTERN

RANGESPHOTOANDREW

CAMPBELL

ADAPTED

BYNIRJA

DESAISCIENCE

on October 18 2018

httpsciencesciencem

agorgD

ownloaded from

commercial farmers that have invested heavily inthemachinery associatedwith chemically intensiveagriculture may not readily switch to agroeco-logical techniques (68 72) however the use ofsome agroecological techniques can be compa-tible with existing infrastructure and can leadto reduced agrochemical use at similar or evenenhanced profits [eg( 73)]A concern is that the use of ldquowildlife-friendlyrdquo

agroecological practices will require more landto be farmed to produce the same amount offood promoting deforestation and harming bio-diversity (74) However a number of diversifiedagroecological farming methods maintain or in-crease yields (table S2) (32 50 73 75ndash78) Forexample techniques such as intercropping covercropping and crop rotation may promote cropyields through a variety of ecologicalmechanisms(23) including complementarity of water andnutrient use (eg different crops access differ-ent soil layers for water and nutrient uptake)facilitation of nutrient uptake [eg intercroppedfaba bean acidifies the soil mobilizing phospho-rus that is taken up by rice (79)] reduction ofpests and diseases [eg pests and diseases spreadmore slowly in spatially or temporally heteroge-neous crop systems and such systems also sup-port predator populations that keep pests incheck (80 81)] and enhancement of soil biotaand fertility (82) By improving soil structure andstability which then enhanceswater infiltrationand retention these techniques also stabilizeyields against annual environmental fluctua-tions and more catastrophic disturbances suchas droughts and hurricanes (32 33)Beyond providing resources and habitats for

agrobiodiversity specific techniques such as agro-forestry and the use of silvopasture hedgerowsflower strips live fences and riparian buffersmay also enhance the connectivity of landscapesand promote the dispersal of various wildlifespecies (16 47 83) Although these structuralfeatures are known to increase the occurrence ofa wide variety of organisms within agriculturallandscapes (43 84) how they affect the dispersalpotential of organisms within diversified agri-cultural lands is poorly understood Nonethelessambitious large-scale connectivity projects suchas the Mesoamerican Biological Corridor project(43) the silvopastoral and rotational grazing proj-ect in the Santa Catarina Atlantic Forest (55)various linkages in Australia (Box 2) and the res-toration of the migratory pathway of the mon-arch butterfly (Danaus plexippus) in the USmidwestern states (85) are underway for agricul-tural lands In the latter case although a daunt-ing amount of restoration would be required tosupport the butterfly it could simultaneously en-hance soybeanpollination improvewater qualityprotect other biodiversity and increase agricul-tural profitability (Fig 5 and table S2) (86 87)Although entrenched policies and the extreme

concentration of agrifood industries favor indus-trialized supply chains andmake transformationto diversified agroecological systems difficult(68 72) reasons for optimism exist Global grass-rootsmovements such as La Via Campesina have

provided technical social and material supportto farmers for the spread of agroecology con-fronted industrial agribusiness and fought toinfluence national and global policies (64) Alter-native agrifood systems and local and regionalinitiatives that provide support for diversifiedagroecological systems are emerging (64 69) In-ternational initiatives supporting agroecologyinclude theUnitedNationsRight toFoodprogramwhich embraces it as a key element for enhancingfood security globally (88) and programs of theFood and Agriculture Organization which hasheld global and regional conferences on agro-ecology and included it in Farmer Field Schoolssince 2014 (68)

Rangelands and forests

Forests in the boreal temperate and tropicalregions make up ~30 of Earthrsquos area (89)whereas rangelands which are defined as havinglt10 tree cover and include grasslands desert

shrublands savannawoodlands alpinemeadowsand areas of tundra grasses and shrubs constitute~44 (90) Grazed by wild and domestic animalsthey vary greatly in productivity Both naturalforests and rangelands have been lost or degradedover the past several hundred years by the in-creased extent and intensity of human use in-cluding timber harvest grazing and conversionto agriculture Forests continue to be lost anddegraded at an alarming rate (62) although for-est regrowth due to rural depopulation is alsooccurring in some areas (20) A recent globalanalysis of sources of tree cover losses showedthat industrial agriculture for commodity cropsis responsible for the permanent conversion of5 million ha of forest per year (27 of losses con-centrated primarily in portions of Latin Americaand Southeast Asia) whereas shifting agriculture(primarily in Africa) and forestry (primarily inNorth America and Europe) cause forest distur-bance or degradation over an equivalent land

Kremen et al Science 362 eaau6020 (2018) 19 October 2018 6 of 9

Monoculture crop Adding prairie strips (10)

8 inchesacre runoff4 tonsacre sediment lost7 lbsacre phosphorus lost35 lbsacre nitrogen lost

42 less runoff95 less soil export89 less phosphorus export84 less nitrogen export

Fig 5 Diversification practices can increase biodiversity The integration of prairie strips into acorn-soy rotation exemplifies how diversification within working lands can substantially increase plantpollinator and bird species richness and abundance by two- to fourfold (as indicated by colors andnumbers of icons respectively) while minimizing externalities and enhancing other ecosystemservices such as pollination for the soy crop (table S2) (86)

RESEARCH | REVIEWILLU

STRATIO

NNIRJA

DESAISCIENCEBASED

ON

STRIPSRESEARCH

TEAM

AND

LEOPOLD

CENTERFO

RSUSTA

INABLE

AGRIC

ULT

URE

on October 18 2018

httpsciencesciencem

agorgD

ownloaded from

area followed by regrowth (62) It is criticaltherefore to cease permanent conversion offorests for commodity cropping and to applyrestorativemanagement approaches inworkingforests and rangelandsSince 1990 many nations have created en-

abling policies and legislation for sustainableforest management (89) Of the 54 of globalforests considered ldquopermanentrdquo (that is expectedto retain forest cover in the long term) 99 ofthese 217 billion ha are covered by such policiesa necessary but not sufficient condition for sus-tainable management Indicators of sustainablemanagement also showpositive temporal trendsbut over smaller areas For example forest cer-tification (table S1) covered 430 million ha by2014 (89) but largely within boreal and temper-ate regions where land-clearing rates are lessacute than those in the tropicsAn array of restorative forest and rangeland

management options exist that are compati-ble with the conservation of working lands (Fig3B and table S2) For forests the adoption ofecosystem-based management approaches hasled to the integration of a greater variety of treespecies and age and size classes including oldgrowth and dead and downed trees and the in-corporation of natural disturbance regimes to sup-port more diverse ecological communities (91)This uneven-aged management style maintainssimilarities betweennatural andmanaged forestscontrasting with even-aged management fromclear-cutting Evidence from silvicultural trials andnatural forests suggests that greater tree diversityalso enhances wood yield quantity and stability(23) In keeping with the ecosystem stewardshipconcept (27) ecosystem-basedmanagement alsoemphasizes collaborative decentralized controland adaptive management as well as landscapeplanning and the designation of corridors to pro-mote wildlife (92) However stakeholders mayreject harvesting practices that negatively af-fect financial returns in the short term Environ-mental outcomes suffered when stakeholdershad stronger oversight of the process than aregulatory authority with political backing (93)supporting the need for public-private part-nerships to achieve biodiversity conservationobjectivesIn rangelands compatible management prac-

tices are exemplified by the dehesa andmontadotraditional pastoral systems in oak savannas ofSpain and Portugal respectively The oak trees(Quercus rotundifolia and Q suber) are prunedto increase the production of acorns to feed topigs and other livestock grown for high-valuemeat products other sustainably harvested pro-ducts include fuelwood and cork from oaks (94)These ecosystems also support endangered spe-cies and high plant and animal diversity rela-tive to other seminatural habitats in EuropeHowever grazing browsing and trampling canlimit oak regeneration thus pasture areas needperiodic temporary protection from livestock topromote oak recruitment and sustainable use(95) In Colombia many ranchers are restoringdegraded agricultural lands by using various

silvopastoral techniques which also enhanceconnectivity in these landscapes (Fig 1)

Freshwater ecosystems

Maintaining stream flows and hydrologic con-nectivity is essential for conserving freshwaterbiodiversity and ecosystems Because of changesin stream flows estimates suggest that up to 75of freshwater fish species are headed for localextinction by 2070 (96) Fresh water also limitsthe production of many natural resources andits quantity and quality are in turn affected bylandscape management Appropriate manage-ment techniques can promote groundwater re-charge and stream flow in working landscapes(table S2) (31 86) of increasing importanceunder drier futures with more extreme precip-itation events (97) Flood plains and associatedriparian zones are particularly critical to conservein working landscapes because they dispropor-tionately support biodiversity and ecosystemprocesses compared with other landscape ele-ments (98) Riparian corridors also provide coolerandmoistermicroclimates than surroundingareasand often span elevational and climatic gradientsthat may permit species to follow their climateenvelopes (99)

Recommendations andconcluding thoughts

Managing the working lands matrix for bio-diversity needs to become amainstream compo-nent of public and private conservation effortscomplementing the more traditional (and essen-tial) focus on increasing the extent and effective-ness of protected areas (16) These restorativeworking lands conservation approaches (tableS2) should be applied to the large land area thatis already used for farming forestry and ranch-ing At the same time we critically need policiesto prevent further conversion and degradation ofwilderness and relatively intact ecosystems (62)To scale up working lands conservation in-

creased support is needed for the voluntary policyand market instruments described in table S1However further adaptation and learning isneeded to improve their efficacy both at theproject level and through evidence-based synthe-ses [eg (100] and to increase adoption ratesby considering an array of social factors (58)Further thesemeasuresmust be complementedby community-driven conservation initiativeswhich by involving young and old in steward-ship communication citizen science and edu-cation can create a shared vision and innovativepractices that result in collective impact Scien-tists can support community-driven conserva-tion and help advance environmental socialmovements by engaging the public listening toalternative ways of knowing and cocreating con-servation management and policy alternativesEspecially important is to create alliances withexisting community actions and socialmovementsthat share common ground such as climate orlocal food movementsUltimately our efforts to protect biodiversity

and sustain resources must be accompanied by

measures to reduce human population and con-sumption while increasing equitable access toresources to achieve sustainability Opportunitiesto stabilize population and consumption existFor example through concerted government in-vestment in voluntary family planning programsenormous progress in reducing total fertilityrates has been made even in poor countries [eg(101] leading to smaller families living betterGlobally a large unmet need for family planningstill exists (101) further investment could helpstabilize the global population at 6 billion peopleby 2100 instead of the 9 to 12 billion projectedwithout intervention (102 103) To reduce con-sumption critical targets include reducing foodwaste and meat consumption (104) and seekingefficiencies in energy and water use that canaccompany urbanization (102) Even with well-structured policies these changes toward lowerhuman population and consumption would taketime thus concerns exist that humanity willdestroy biodiversity and natural resources beforeachieving a more sustainable human population(102) Conservation in working landscapes canhelpmaintain all species including people as westrive to achieve a planet where a smaller humanpopulation lives better and more equitably withand because of wild nature

REFERENCES AND NOTES

1 J L Payne A M Bush N A Heim M L KnopeD J McCauley Ecological selectivity of the emerging massextinction in the oceans Science 353 1284ndash1286 (2016)doi 101126scienceaaf2416 pmid 27629258

2 G Ceballos P R Ehrlich R Dirzo Biological annihilation viathe ongoing sixth mass extinction signaled by vertebratepopulation losses and declines Proc Natl Acad Sci USA114 E6089ndashE6096 (2017) pmid 28696295

3 G Ceballos et al Accelerated modern human-induced specieslosses Entering the sixth mass extinction Sci Adv 1 e1400253(2015) doi 101126sciadv1400253 pmid 26601195

4 C A Hallmann et al More than 75 percent decline over27 years in total flying insect biomass in protected areasPLOS ONE 12 e0185809 (2017) doi 101371journalpone0185809 pmid 29045418

5 T Newbold et al Global effects of land use on localterrestrial biodiversity Nature 520 45ndash50 (2015)doi 101038nature14324 pmid 25832402

6 E O Wilson Half Earth (Liveright Publishing Company 2016)7 P R Elsen W B Monahan A M Merenlender Global

patterns of protection of elevational gradients in mountainranges Proc Natl Acad Sci USA 115 6004ndash6009 (2018)doi 101073pnas1720141115 pmid 29784825

8 R M Pringle Upgrading protected areas to conserve wildbiodiversity Nature 546 91ndash99 (2017) doi 101038nature22902 pmid 28569807

9 J M Halley N Monokrousos A D Mazaris W D NewmarkD Vokou Dynamics of extinction debt across five taxonomicgroups Nat Commun 7 12283 (2016) doi 101038ncomms12283 pmid 27452815

10 W F Laurance et al Averting biodiversity collapse in tropicalforest protected areas Nature 489 290ndash294 (2012)doi 101038nature11318 pmid 22832582

11 M C Urban Climate change Accelerating extinction riskfrom climate change Science 348 571ndash573 (2015)doi 101126scienceaaa4984 pmid 25931559

12 S R Loarie et al The velocity of climate change Nature 4621052ndash1055 (2009) doi 101038nature08649pmid 20033047

13 L O Frishkoff et al Loss of avian phylogeneticdiversity in neotropical agricultural systems Science345 1343ndash1346 (2014) doi 101126science1254610pmid 25214627

14 G Chapron et al Recovery of large carnivores in Europersquosmodern human-dominated landscapes Science 346 1517ndash1519(2014) doi 101126science1257553 pmid 25525247

Kremen et al Science 362 eaau6020 (2018) 19 October 2018 7 of 9

RESEARCH | REVIEWon O

ctober 18 2018

httpsciencesciencemagorg

Dow

nloaded from

15 M D Reynolds et al Dynamic conservation for migratoryspecies Sci Adv 3 e1700707 (2017) doi 101126sciadv1700707 pmid 28845449

16 C Kremen Reframing the land-sparingland-sharingdebate for biodiversity conservation Ann NY Acad Sci1355 52ndash76 (2015) doi 101111nyas12845pmid 26213864

17 C D Mendenhall A Shields-Estrada A J KrishnaswamiG C Daily Quantifying and sustaining biodiversity in tropicalagricultural landscapes Proc Natl Acad Sci USA 11314544ndash14551 (2016) doi 101073pnas1604981113pmid 27791070

18 C D Mendenhall D S Karp C F J Meyer E A HadlyG C Daily Predicting biodiversity change and avertingcollapse in agricultural landscapes Nature 509 213ndash217(2014) doi 101038nature13139 pmid 24739971

19 C M Kennedy E H C Grant M C Neel W F FaganP P Marra Landscape matrix mediates occupancy dynamicsof Neotropical avian insectivores Ecol Appl 21 1837ndash1850(2011) doi 10189010-10441 pmid 21830722

20 J A Foley et al Global consequences of land use Science309 570ndash574 (2005) doi 101126science1111772pmid 16040698

21 N Ramankutty et al Trends in global agricultural land useImplications for environmental health and food securityAnnu Rev Plant Biol 69 789ndash815 (2018) doi 101146annurev-arplant-042817-040256 pmid 29489395

22 F Isbell et al Linking the influence and dependenceof people on biodiversity across scales Nature 546 65ndash72(2017) doi 101038nature22899 pmid 28569811

23 F Isbell et al Benefits of increasing plant diversity insustainable agroecosystems J Ecol 105 871ndash879 (2017)doi 1011111365-274512789

24 D J Eastburn A T OrsquoGeen K W Tate L M Roche Multipleecosystem services in a working landscape PLOS ONE 12e0166595 (2017) doi 101371journalpone0166595pmid 28301475

25 D A Driscoll S C Banks P S Barton D B LindenmayerA L Smith Conceptual domain of the matrix in fragmentedlandscapes Trends Ecol Evol 28 605ndash613 (2013)doi 101016jtree201306010 pmid 23883740

26 W F Fagan E E Holmes Quantifying the extinction vortexEcol Lett 9 51ndash60 (2006) pmid 16958868

27 F S Chapin 3rd et al Ecosystem stewardship Sustainabilitystrategies for a rapidly changing planet Trends Ecol Evol 25241ndash249 (2010) doi 101016jtree200910008pmid 19923035

28 F Isbell et al High plant diversity is needed to maintainecosystem services Nature 477 199ndash202 (2011)doi 101038nature10282 pmid 21832994

29 M G E Mitchell E M Bennett A Gonzalez Linkinglandscape connectivity and ecosystem service provisionCurrent knowledge and research gaps Ecosystems 16894ndash908 (2013) doi 101007s10021-013-9647-2

30 M Loreau N Mouquet A Gonzalez Biodiversity as spatialinsurance in heterogeneous landscapes Proc Natl Acad SciUSA 100 12765ndash12770 (2003) doi 101073pnas2235465100 pmid 14569008

31 C Kremen A Miles Ecosystem services in biologicallydiversified versus conventional farming systems Benefitsexternalities and trade-offs Ecol Soc 17 40 (2012)doi 105751ES-05035-170440

32 A C M Gaudin et al Increasing crop diversity mitigatesweather variations and improves yield stability PLOS ONE 10e0113261 (2015) doi 101371journalpone0113261pmid 25658914

33 S M Philpott B B Lin S Jha S J Brines A multi-scaleassessment of hurricane impacts on agricultural landscapesbased on land use and topographic features AgricEcosyst Environ 128 12ndash20 (2008) doi 101016jagee200804016

34 L C Ponisio et al Diversification practices reduce organic toconventional yield gap Proc R Soc London Ser B 28220141396 (2015) doi 101098rspb20141396pmid 25621333

35 R J Hobbs et al Managing the whole landscape Historicalhybrid and novel ecosystems Front Ecol Environ 12557ndash564 (2014) doi 101890130300

36 N Estrada-Carmona A K Hart F A J DeClerckC A Harvey J C Milder Integrated landscape managementfor agriculture rural livelihoods and ecosystem conservationAn assessment of experience from Latin America andthe Caribbean Landsc Urban Plan 129 1ndash11 (2014)doi 101016jlandurbplan201405001

37 H Gu S M Subramanian Drivers of change insocio-ecological production landscapes Implications forbetter management Ecol Soc 19 41 (2014) doi 105751ES-06283-190141

38 M K Anderson Tending the Wild Native American Knowledgeand the Management of Californiarsquos Natural Resources(Univ of California Press 2005)

39 E M Bignal D I McCracken The nature conservation valueof European traditional farming systems Environ Rev 8149ndash171 (2000) doi 101139a00-009

40 S K Lowder J Skoet T Raney The number size anddistribution of farms smallholder farms and family farmsworldwide World Dev 87 16ndash29 (2016) doi 101016jworlddev201510041

41 C L Gray et al Local biodiversity is higher inside thanoutside terrestrial protected areas worldwide Nat Commun7 12306 (2016) doi 101038ncomms12306pmid 27465407

42 C L Shafer Cautionary thoughts on IUCN protected areamanagement categories V-VI Global Ecol Conserv 3331ndash348 (2015) doi 101016jgecco201412007

43 F A J DeClerck et al Biodiversity conservation inhuman-modified landscapes of Mesoamerica Past presentand future Biol Conserv 143 2301ndash2313 (2010)doi 101016jbiocon201003026

44 C A Harvey et al Patterns of animal diversityin different forms of tree cover in agriculturallandscapes Ecol Appl 16 1986ndash1999 (2006)doi 1018901051-0761(2006)016[1986POADID]20CO2pmid 17069389

45 Ccedil H Şekercioğlu et al Tropical countryside ripariancorridors provide critical habitat and connectivity forseed-dispersing forest birds in a fragmented landscapeJ Ornithol 156 (Suppl 1) 343ndash353 (2015) doi 101007s10336-015-1299-x

46 C A Harvey et al Contribution of live fences to theecological integrity of agricultural landscapesAgric Ecosyst Environ 111 200ndash230 (2005) doi 101016jagee200506011

47 A Medina C A Harvey D S Merlo S Viacutelchez B HernaacutendezBat diversity and movement in an agricultural landscape inMatiguaacutes Nicaragua Biotropica 39 120ndash128 (2007)doi 101111j1744-7429200600240x

48 K Williams-Guillen C McCann J C Martinez SanchezF Koontz Resource availability and habitat use by mantledhowling monkeys in a Nicaraguan coffee plantation Canagroforests serve as core habitat for a forest mammalAnim Conserv 9 331ndash338 (2006) doi 101111j1469-1795200600042x

49 J Avelino A Romero-Gurdiaacuten H F Cruz-CuellarF A J Declerck Landscape context and scale differentiallyimpact coffee leaf rust coffee berry borer and coffeeroot-knot nematodes Ecol Appl 22 584ndash596 (2012)doi 10189011-08691 pmid 22611856

50 D S Karp et al Forest bolsters bird abundance pest controland coffee yield Ecol Lett 16 1339ndash1347 (2013)doi 101111ele12173 pmid 23981013

51 C A Harvey et al Climate-smart landscapes Opportunitiesand challenges for integrating adaptation and mitigation intropical agriculture Conserv Lett 7 77ndash90 (2014)doi 101111conl12066

52 A T H Keeley et al Making habitat connectivity areality Conserv Biol (2018) doi 101111cobi13158pmid 29920775

53 W C Morse et al Consequences of environmental servicepayments for forest retention and recruitment in a Costa Ricanbiological corridor Ecol Soc 14 23 (2009) doi 105751ES-02688-140123

54 J Owley D Takacs ldquoFlexible Conservation in UncertainTimesrdquo in Contemporary Issues in Climate Change Law andPolicy Essays Inspired by the IPCC (UC Hastings ResearchPaper 180 Univ at Buffalo School of Law 2016)pp 65ndash102 httpspapersssrncomsol3paperscfmabstract_id=2700658

55 J Alvez A L Schmitt Filho J Farley G Alarcon A C FantiniThe potential for agroecosystems to restore ecologicalcorridors and sustain farmer livelihoods Evidence fromBrazil Ecol Restor 30 288ndash290 (2012) doi 103368er304288

56 C M Kennedy et al Bigger is better Improved natureconservation and economic returns from landscape-levelmitigation Sci Adv 2 e1501021 (2016) doi 101126sciadv1501021 pmid 27419225

57 I Scoones P Newell M Leach in The Politics of GreenTransformations I Scoones M Leach P Newell Eds(Earthscan from Routledge 2015) pp 1ndash24

58 M B Mascia M Mills When conservation goes viralThe diffusion of innovative biodiversity conservationpolicies and practices Conserv Lett 11 e12442 (2018)doi 101111conl12442

59 D F Doak V J Bakker B E Goldstein B Hale Whatis the future of conservation Trends Ecol Evol 29 77ndash81(2014) doi 101016jtree201310013 pmid 24332874

60 K Hamrick ldquoState of private investment in conservation2016 A landscape assessment of an emerging marketrdquo(Forest Trends 2016)

61 R Chaplin-Kramer et al Ecosystem service information tobenefit sustainability standards for commodity supply chainsAnn NY Acad Sci 1355 77ndash97 (2015) doi 101111nyas12961 pmid 26555859

62 P G Curtis C M Slay N L Harris A TyukavinaM C Hansen Classifying drivers of global forest lossScience 361 1108ndash1111 (2018) doi 101126scienceaau3445 pmid 30213911

63 D Pannell et al Understanding and promoting adoption ofconservation practices by rural landholders Aust J ExpAgric 46 1407ndash1424 (2006) doi 101071EA05037

64 M Leach I Scoones in The Politics of GreenTransformations I Scoones P Newell M Leach Eds(Earthscan from Routledge 2015) pp 119ndash133

65 D R Cameron D C Marvin J M Remucal M C PasseroEcosystem management and land conservation cansubstantially contribute to Californiarsquos climate mitigationgoals Proc Natl Acad Sci USA 114 12833ndash12838 (2017)doi 101073pnas1707811114 pmid 29133408

66 N Ramankutty A T Evan C Monfreda J A Foley Farmingthe planet 1 Geographic distribution of global agriculturallands in the year 2000 Global Biogeochem Cycles 22GB1003 (2008) doi 1010292007GB002952

67 M A Altieri The ecological role of biodiversity inagroecosystems Agric Ecosyst Environ 74 19ndash31 (1999)doi 101016S0167-8809(99)00028-6

68 International Panel of Experts on Sustainable Food Systems(IPES-Food) ldquoFrom uniformity to diversity A paradigmshift from industrial agriculture to diversified agroecologicalsystemsrdquo (IPES-Food 2016) httpwwwipes-foodorgimagesReportsUniformityToDiversity_FullReportpdf

69 C Kremen A Iles C M Bacon Diversified farming systemsAn agro-ecological systems-based alternative to modernindustrial agriculture Ecol Soc 17 44 (2012) doi 105751ES-05103-170444

70 V Ricciardi N Ramankutty Z Mehrabi L JarvisB Chookolingo How much of the worldrsquos food dosmallholders produce Global Food Sec 17 64ndash72 (2018)doi 101016jgfs201805002

71 M Montenegro Banking on wild relatives to feed the worldGastron J Crit Food Stud 16 1ndash8 (2016) doi 101525gfc20161611

72 A Iles R Marsh Nurturing diversified farming systemsin industrialized countries How public policy cancontribute Ecol Soc 17 42 (2012) doi 105751ES-05041-170442

73 A S Davis J D Hill C A Chase A M JohannsM Liebman Increasing cropping system diversity balancesproductivity profitability and environmental health PLOSONE 7 e47149 (2012) doi 101371journalpone0047149pmid 23071739

74 B Phalan M Onial A Balmford R E Green Reconciling foodproduction and biodiversity conservation Land sharing andland sparing compared Science 333 1289ndash1291 (2011)doi 101126science1208742 pmid 21885781

75 K Garbach et al Examining multi-functionality for crop yieldand ecosystem services in five systems of agroecologicalintensification Int J Agric Sustain 15 11ndash28 (2017)doi 1010801473590320161174810

76 J Pretty Z P Bharucha Sustainable intensification inagricultural systems Ann Bot 114 1571ndash1596 (2014)doi 101093aobmcu205 pmid 25351192

77 G M Gurr et al Multi-country evidence that crop diversificationpromotes ecological intensification of agricultureNat Plants 2 16014 (2016) doi 101038nplants201614pmid 27249349

78 R F Pywell et al Wildlife-friendly farming increases cropyield Evidence for ecological intensification Proc R SocLondon Ser B 282 20151740 (2015) doi 101098rspb20151740 pmid 26423846

Kremen et al Science 362 eaau6020 (2018) 19 October 2018 8 of 9

RESEARCH | REVIEWon O

ctober 18 2018

httpsciencesciencemagorg

Dow

nloaded from

79 L Li et al Diversity enhances agricultural productivity viarhizosphere phosphorus facilitation on phosphorus-deficientsoils Proc Natl Acad Sci USA 104 11192ndash11196 (2007)doi 101073pnas0704591104 pmid 17592130

80 D K Letourneau et al Does plant diversity benefitagroecosystems A synthetic review Ecol Appl 21 9ndash21(2011) doi 10189009-20261 pmid 21516884

81 Y Zhu et al Genetic diversity and disease control in riceNature 406 718ndash722 (2000) doi 10103835021046pmid 10963595

82 S F Bender C Wagg M G A van der Heijden Anunderground revolution Biodiversity and soil ecologicalengineering for agricultural sustainability Trends Ecol Evol 31440ndash452 (2016) doi 101016jtree201602016 pmid26993667

83 I Perfecto J Vandermeer A Wright Naturersquos Matrix LinkingAgriculture Conservation and Food Sovereignty (Earthscan 2009)

84 J A Hilty A M Merenlender Use of riparian corridors andvineyards by mammalian predators in northern CaliforniaConserv Biol 18 126ndash135 (2004) doi 101111j1523-1739200400225x

85 W E Thogmartin et al Restoring monarch butterfly habitatin the Midwestern US lsquoAll hands on deckrsquo Environ Res Lett12 074005 (2017) doi 1010881748-9326aa7637

86 L A Schulte et al Prairie strips improve biodiversityand the delivery of multiple ecosystem services fromcorn-soybean croplands Proc Natl Acad Sci USA 11411247ndash11252 (2017) doi 101073pnas1620229114pmid 28973922

87 E Brandes et al Subfield profitability analysis reveals aneconomic case for cropland diversification Environ Res Lett11 014009 (2016) doi 1010881748-9326111014009

88 O De Schutter ldquoAgroecology and the right to foodrdquo reportpresented at the 16th Session of the United NationsHuman Rights Council [AHRC1649] Geneva Switzerland8 March 2011

89 K G MacDicken et al Global progress toward sustainableforest management For Ecol Manage 352 47ndash56 (2015)doi 101016jforeco201502005

90 H G Lund Accounting for the worldrsquos rangelandsSoc Range Manage 29 3ndash10 (2007)

91 S Gauthier M-A Vaillancourt D Kneeshaw P DrapeauL De Grandpreacute Y Claveau D Pareacute in Ecosystem Managementin the Boreal Forest S Gauthier M-A Vaillancourt A LeducL De Grandpreacute D Kneeshaw H Morin P DrapeauY Bergeron Eds (Univ du Queacutebec 2009) pp 13ndash38

92 M Mangel et al Principles for the conservation of wild livingresources Ecol Appl 6 338ndash362 (1996) doi 1023072269369

93 J A Layzer Natural Experiments Ecosystem-BasedManagement and the Environment (MIT 2008)

94 P Campos L Huntsinger J L Oviedo P F Starrs M DiazR B Standiford G Montero Eds Mediterranean OakWoodland Working Landscapes Dehesas of Spain and Ranchlandsof California (Springer Science+Business Media 2013)

95 J A Ramiacuterez M Diacuteaz The role of temporal shrubencroachment for the maintenance of Spanish holm oakQuercus ilex dehesas For Ecol Manage 255 1976ndash1983(2008) doi 101016jforeco200712019

96 M A Xenopoulos et al Scenarios of freshwater fish extinctionsfrom climate change and water withdrawal Global Change Biol11 1557ndash1564 (2005) doi 101111j1365-24862005001008x

97 S D Polade A Gershunov D R Cayan M D DettingerD W Pierce Precipitation in a warming world Assessingprojected hydro-climate changes in California and otherMediterranean climate regions Sci Rep 7 10783 (2017)doi 101038s41598-017-11285-y pmid 28883636

98 F R Hauer et al Gravel-bed river floodplains are theecological nexus of glaciated mountain landscapes Sci Adv2 e1600026 (2016) doi 101126sciadv1600026pmid 27386570

99 A T Keeley et al New concepts models and assessmentsof climate-wise connectivity Environ Res Lett 13 073002(2018) doi 1010881748-9326aacb85

100 W Sutherland L V Dicks N Ockendon R Smith Eds WhatWorks in Conservation (Open Book ed 2 2017) vol 2

101 J Bongaarts S W Sinding A response to critics of familyplanning programs Int Perspect Sex Reprod Health 3539ndash44 (2009) doi 1013633503909 pmid 19465347

102 E W Sanderson J Walston J G Robinson From bottleneckto breakthrough Urbanization and the future of biodiversityconservation Bioscience 68 412ndash426 (2018) doi 101093bioscibiy039 pmid 29867252

103 C J Bradshaw B W Brook Human population reduction isnot a quick fix for environmental problems Proc NatlAcad Sci USA 111 16610ndash16615 (2014) doi 101073pnas1410465111 pmid 25349398

104 J Poore T Nemecek Reducing foodrsquos environmentalimpacts through producers and consumers Science360 987ndash992 (2018) doi 101126scienceaaq0216pmid 29853680

105 Great Eastern Ranges wwwgerorgau106 V Athreya M Odden J D C Linnell J Krishnaswamy

U Karanth Big cats in our backyards Persistence of largecarnivores in a human dominated landscape in IndiaPLOS ONE 8 e57872 (2013) doi 101371journalpone0057872pmid 23483933

ACKNOWLEDGMENTS

We appreciate the constructive input of D Ackerly B BrunnerA Campbell F DeClerck and A Knight Competing interests Theauthors declare no competing interests

SUPPLEMENTARY MATERIALS

wwwsciencemagorgcontent3626412eaau6020supplDC1Tables S1 and S2References (107ndash158)

101126scienceaau6020

Kremen et al Science 362 eaau6020 (2018) 19 October 2018 9 of 9

RESEARCH | REVIEWon O

ctober 18 2018

httpsciencesciencemagorg

Dow

nloaded from

Landscapes that work for biodiversity and peopleC Kremen and A M Merenlender

DOI 101126scienceaau6020 (6412) eaau6020362Science

this issue p eaau6020Sciencejust for ecosystem services but also for maintenance and persistence of nonhuman speciesmost human-modified lands as working landscapes These can provide for human needs and maintain biodiversity nothave to be a lost cause Kremen and Merenlender review how biodiversity-based techniques can be used to manage and preserved However this still leaves vast regions of the world unprotected and modified Such landscapes do notnonhuman species This is clearly unsustainable and the amount of land we protect for nature needs to be increased

As the human population has grown we have taken and modified more and more land leaving less and less forA nature-friendly matrix

ARTICLE TOOLS httpsciencesciencemagorgcontent3626412eaau6020

MATERIALSSUPPLEMENTARY httpsciencesciencemagorgcontentsuppl201810173626412eaau6020DC1

CONTENTRELATED httpsciencesciencemagorgcontentsci3626412287full

REFERENCES

httpsciencesciencemagorgcontent3626412eaau6020BIBLThis article cites 142 articles 29 of which you can access for free

PERMISSIONS httpwwwsciencemagorghelpreprints-and-permissions

Terms of ServiceUse of this article is subject to the

is a registered trademark of AAASSciencelicensee American Association for the Advancement of Science No claim to original US Government Works The title Science 1200 New York Avenue NW Washington DC 20005 2017 copy The Authors some rights reserved exclusive

(print ISSN 0036-8075 online ISSN 1095-9203) is published by the American Association for the Advancement ofScience

on October 18 2018

httpsciencesciencem

agorgD

ownloaded from