Content uploaded by Adina Merenlender
Author content
All content in this area was uploaded by Adina Merenlender on Mar 12, 2020
Content may be subject to copyright.
REVIEW SUMMARY
◥
CONSERVATION
Landscapes that work for biodiversity
and people
C. Kremen*and A. M. Merenlender
BACKGROUND: Biodiversity is under siege,
with greatly enhanced rates of local and global
extinction and the decline of once-abundant
species. Current rates of human-induced climate
change and land use forecast the Anthropocene
as one of the most devastating epochs for life
on earth. How do we handle the Anthropocene’s
triple challenge of preventing biodiversity loss,
mitigating and adapting to climate change, and
sustainably providing resources for a growing
human population? The answer is in how we
manage Earth’s“working lands”; that is, farms,
forests, and rangelands. These lands must be
managed both to complement the biodiversity
conservation goals of protected ar eas and to
maintain the diverse communities of orga-
nisms, from microbes to mammals, that con-
tribute to producing food, materials, clean
water, and healthy soils; sequest eri ng gree n-
house gases; and buffering extreme weather
events, functions that are essential for all life
on Earth.
ADVANCES: Protected areas are the corner-
stone of biodiversity conservation. Although
the total area of protected regions needs to be
increased, parks will nonetheless continue to
lose species if these areas are isolated from one
another by inhospitable land uses and are
faced with a rapidly changing climate. Further,
many species, such as those that migrate, re-
main unprotected as they occupy lands outside
of parks for all or portions of their life cycles.
Lastly, protected-area effectiveness is greatly
influenced by surrounding land management.
“Working lands conservation”aims to sup-
port biodiversity while providing goods and
services for humanity over the long term, assur-
ing sustainability and resilience. By manag-
ing lands surrounding parks favorably, working
lands can buffer protected areas from threats
and connect them to one another. This ap-
proach complements protected areas by pro-
viding accessory habitats and resources for
some species while facilitating dispersal and
climate change adaptation for others. Further,
by maintaining the biodiversity that supplies
critical ecosystem services within working
lands, these approaches ensure that the pro-
duction of food, fiber, fuel, and timber can be
sustained over the long run and be more resil-
ient to extreme events, such as floods, dro ughts,
hurricanes, and pest and disease outbreaks, which
are becoming more frequent with climate change.
A variety of biodiversity-based land management
techniques can be used in working lands, includ-
ing agroforestry, silvopasture, diversified farming,
and ecosystem-based forest management, to en-
sure sustainable production of food and fiber.
OUTLOOK: The underlying principle of
biodiversity-based management of working
lands has been practiced since ancient times.
Today, these systems have largely been replaced
by unsustainable resource extraction, rather
than serving as models that could be adapted
to modern conditions. Although various reg-
ulatory, voluntary, and financial tools exist to
promote sustainable land management, many
barriers prevent individuals, communities, and
corporations from adopting biodiversity-based
practices, including deeply entrenched policy
and market conditions that favor i ndustrialized
or extractive models of land use. Thus, uptake
of these approaches has
been patchy and slow and
is not yet sufficient to cre-
ate change at the tempo-
ral and spatial scales needed
to face the triple Anthro-
pocene threat.
Biodiversity-based land management prac-
tices are knowledge- rather than technology-
intensive. They are well adapted to empower
local communities to manage their natural
resources. One of the most exciting emerging
trends is community-driven initiatives to man-
age working landscapes for conservation and
sustainability. By linking up through grass-
roots organizations, social movements, and
public-private partnerships, these initiatives
can scale up to create collective impact and can
demand changes in government policies to
facilitate the conservation of working lands.
Scientists and conservation practitioners can
support these initiatives by engaging with the
public, listening to alternative ways of knowing,
and cocreating landscapes that work for bio-
diversity and people.▪
RESEARCH
Kremen et al., Science 362, 304 (2018) 19 October 2018 1of1
TOMORROW’SEARTH
Read more articles online
at scim.ag/TomorrowsEarth
The list of author affiliations is available in the full article online.
*Corresponding author. Email: ckremen@berkeley.edu
Cite this article as C. Kremen and A. M. Merenlender, Science
362, eaau6020 (2018). DOI: 10.1126/science.aau6020
Strawberry production in
Central Coast, California. On
the left, a homogeneous land-
scape of strawberry mono-
culture, including organic fields,
supports fewer wild species
then a diversified, organic farm
(right) in the same region, which
includes a small field of straw-
berry, surrounded by orchards,
hedgerows, diverse vegetable
crops, and natural habitats. The
monoculture landscape creates
barriers to wildlife dispersal,
whereas the diversified land-
scape is more permeable.
PHOTO: C. KREMEN
ON OUR WEBSITE
◥
Read the full article
at http://dx.doi.
org/10.1126/
science.aau6020
..................................................
on October 18, 2018 http://science.sciencemag.org/Downloaded from
REVIEW
◥
CONSERVATION
Landscapes that work for biodiversity
and people
C. Kremen*and A. M. Merenlender
How can we manage farmlands, forests, and rangelands to respond to the triple challenge
of the Anthropocene—biodiversity loss, climate change, and unsustainable land use?
When managed by using biodiversity-based techniques such as agroforestry, silvopasture,
diversified farming, and ecosystem-based forest management, these socioeconomic systems
can help maintain biodiversity and provide habitat connectivity, thereby complementing
protected areas and providing greater resilience to climate change. Simultaneously, the
use of these management techniques can improve yields and profitability more sustainably,
enhancing livelihoods and food security. This approach to “working lands conservation”can
create landscapes that work for nature and people. However, many socioeconomic challenges
impede the uptake of biodiversity-based land management practices. Although improving
voluntary incentives, market instruments, environmental regulations, and governance is
essential to support working lands conservation, it is community action, social movements,
and broad coalitions among citizens, businesses, nonprofits, and government agencies that
have the power to transform how we manage land and protect the environment.
Biodiversity, the product of 3.8 billion years
of evolution, is under siege. Not only are
both marine and terrestrial species expe-
riencing accelerated rates of local and global
extinction (1–3), but even common species
are declining (2,4,5). This alarming situation
has prompted a strong call for increasing the
number (6,7) and effectiveness (8) of protected
areas, the principal method for combatting spe-
cies loss. Though such protections are essential, we
cannot rely on protected areas alone to preserve
species. As protected areas become increasingly
isolated because of habitat loss and degradation,
much research has revealedthattheywilllosespe-
cies over time (9). Further, many critical threats to
species do not respect protected-area boundaries
(10), including climate change, which both exac-
erbates species losses (11) and threatens to alter
the biomes of many currently protected regions
entirely (12).
More hopefully, recent studies show that some
human-dominated landscapes can support much
more biodiversity than previously recognized
(13–17), suggesting a complementary path for-
ward. Specifically, when these areas, generally
referred to as the “matrix,”represent a high-
quality mosaic of land uses, they can play a crit-
ical role in sustaining biodiversity, both in situ
and by promoting species dispersal among pro-
tected areas and remnant habitats and along
migratory routes (Fig. 1) (15,18,19). Of course,
human survival also depends on the long-term
capacity of this matrix of “working lands,”in-
cluding rangelands, forests, and farms, to pro-
duce food, water, fiber, fuel, and forest products.
All too often, however, these goods are produced
at severe environmental cost, including habitat
degradation, toxic contamination, and depletion
of water quantity and quality, leading to ecologi-
cal collapse, local extinctions, and the creation of
unproductive wastelands (20,21). We argue that,
instead, working lands can be used to support
high levels of biodiversity while satisfying human
needs in a sustainable way. Because rangelands,
forests, and cultivated lands collectively occupy
~80% of terrestrial area (21), the potential for con-
servationinsuchlandsisenormous.
Critical ecosystem functions and services are
provided by a suite of diverse organisms, from
microbes to mammals, and thus maintenance
of these organisms is necessary for long-term
and sustainable productivity of working lands
(22,23). Hence, managing the matrix to main-
tain biodiversity is not only necessary for species
conservation but also essential for sustainable pro-
duction. Biodiversity-based production systems,
including agroecological farming or ecosystem-
based forest management, are often perceived as
unproductive, an incorrect viewpoint that im-
pedes the public investment needed to develop
and promote these methods. Here, we describe
managingthematrixjointlyandsustainablyfor
biodiversity and people through “working lands
conservation”and ask what strategies can be used
to strengthen and scale up this approach as
rapidlyaspossibletohelpcombatthetriple
Anthropocene threats of biodiversity loss, cli-
mate change, and unsustainable land use.
Working lands conservation defined
Although the term “working lands conservation”
is already used in policy statements and in guid-
ance for conservation programs [e.g., (24)], the
concept has yet to be formally defined and risks
being misapplied. We define it at the landscape
scale (Box 1).
To avoid mass extinction and ecosystem col-
lapse, we must integrate biodiversity conserva-
tion into the landscapes we use and not simply
relegate nature to a limited number of protected
areas that are doomed if left as isolated habitat
islands within biological deserts. Working lands
can provide food, breeding sites, and shelter for
a myriad of species while maintaining abiotic
conditions, including temperature, light, wind,
water, fire, and other disturbance processes,
within required ranges. They can facilitate func-
tional connectivity—that is, the movement of orga-
nisms across the landscape and among habitat
patches that promotes population persistence by
allowing 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 the
ecosystem services, such as pollination, pest
control, and nutrient cycling, that underlie that
production. Maintaining these services requires
supporting the underlying populations of service-
providing organisms. Within each service, a greater
diversity of service providers often enhances the
level and/or quality of services and reduces un-
certainty in service delivery (22), because different
species respond differentially to environmental
change (27,28). Maintaining connectivity is also
important, both to support flows of ecosystem
service providers and/or materials (e.g., pollination
requires animal vectors to move pollen between
flowers; water purification requires water to flow
through vegetation) (29) and to enhance meta-
community persistence of service-providing orga-
nisms to sustain ecosystem functions and services
over space and time (22,30).
RESEARCH
Kremen et al., Science 362, eaau6020 (2018) 19 October 2018 1of9
Department of Environmental Science, Policy and
Management, University of California, Berkeley, Berkeley,
CA 94720, USA.
*Corresponding author. Email: ckremen@berkeley.edu
Box 1. Definition of working lands conservation.
Definition: Conservation in working landscapes maintains biodiversity, provides goods and services
for humanity, and supports the abiotic conditions necessary for sustainability and resilience.
These socioecological systems both support biodiversity by providing critical resources and
rely on biodiversity (specifically, ecosystem service providers) for sustainable production of
food, water, fiber, fuel, and forest products. These landscapes also enhance connectivity to
promote the movement of organisms, natural processes, and ecosystem services.
Working lands conservation emphasizes the critical role of managing the matrix for species
conservation to complement protected areas.
on October 18, 2018 http://science.sciencemag.org/Downloaded from
Ensuring the sustainability of production re-
quires balancing across provisioning, regulat-
ing, and supporting services; in other words,
seeking multifunctionality and stability rather
than maximal production. For example, conven-
tional (chemically intensive) monoculture agri-
culture produces high yields but often at the
expense of water quality, climate regulation, and
soil health (Fig. 2A) (20)andcansufferproduc-
tion collapse in response to periodic extreme
weather, pests, and diseases (31–33). Although
transforming to a more sustainable system may
reduce average yields somewhat [e.g., (34)], by
relying on ecosystem services produced on the
farm and in the surrounding landscape, a sus-
tainable system is both multifunctional and more
resilient to change (20,31)(Fig.2C).
Working landscapes often comprise hetero-
geneous patch types, including novel commu-
nities made up of mixtures of native and nonnative
species, as well as remnants of natural or semi-
natural habitats whose composition is more simi-
lar to that of a historical ecological community
(35). Although management goals likely will dif-
fer among patch types, both individual patches
and the whole landscape should be managed
for sustainability. For example, patches whose
communities are far from historical could be man-
aged principally for crops (a provisioning service)
by using sustainable agricultural practices to min-
imize negative effects on biodiversity and ecosys-
tem services on and off site. Remnant patches
could be retained as stepping-stone habitats to
support species dispersal and provide regulating
services such as pollination (29,31). Maintaining
mosaic landscapes composed of different patch
types provides opportunities to maximize diver-
sity, resilience, and multifunctionality. Radar dia-
grams reveal likely trade-offs and sustainability
within and across patches (Fig. 2B), as well as
multifunctionality 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, biodiversity
or ecosystem conservation, and rural livelihoods
and are being implemented by governments and
nongovernmental organizations in Latin America
and Africa (36). The ecosystem stewardship con-
cept focuses on the need to sustain Earth’s capac-
ity to provide ecosystem services and support
socioecological resilience under conditions of
uncertainty and change (27). The socioecological
production landscape of the Japan Satoyama
Satoumi Assessment refers to dynamic landscape
mosaics that have been shaped over time by the
interactions between people and nature in ways
that jointly support biodiversity and human well-
being (37). These concepts also emphasize critical
social components, such as involving multiple
stakeholders at the landscape scale, community
participation, intersectoral coordination, flexible
and adaptive governance systems, social learn-
ing, and adaptive management, which are nec-
essary for successful conservation of working
landscapes.
The underlying principle of maintaining eco-
logical diversity inherent to these approaches
has been practiced since ancient times. Some of
these management systems, such as indigenous
use of fire, weeding, pruning, and the seed dis-
persal that shaped Californian ecosystems (38), no
longer exist in their original form, whereas others,
such as regional pastoral and high-mountain farm-
ing systems in Europe (39), persist in some areas.
By creating highly simplified and intensified pro-
duction systems (21,40), from corn and soy in
U.S. midwestern states to palm oilplantations in
southeast Asia and vineyards in Chile, we have
abandoned this critical sustainability principle
across much of Earth’scultivatedlandscapes.
However, it is a fallacy that such systems will
ultimatelysparemorelandfornatureconser-
vation or feed the world indefinitely; rather, we
need to find ways to allow biodiversity-based
production methods to figure much more prom-
inently in local, regional, and global markets (16).
Working lands conservation as a
complement to protected areas
Given the dire situation facing many species and
the expectation of further species losses and shifts
in ecosystem composition due to climate change
(2,4,11), ceasing further habitat conversion
completely and protecting large regions of Earth
effectively are critical necessities for conservation
(6–8), although just how much should be pro-
tected is highly debated (41). [By “protected area,”
we refer to parks whose primary function is to
conserve biodiversity and wilderness (Interna-
tional Union for Conservation of Nature and
Natural Resources categories I to IV, constituting
6.75% of terrestrial area) (42), in contrast to areas
blending conservation and livelihood objectives
(categories V to VI, constituting 8.65%).] How-
ever, the protected-area strategy alone will not
be successful without complementary working
lands conservation in the surrounding landscapes.
First, even the largest protected areas will lose
species over the long term (9) unless surrounding
landscapes can be managed to provide connec-
tivity among parks. Further, less than 10% of pro-
tected areas are expected to represent current
climatic conditions within 100 years, increasing
the criticality of matrix connectivity to permit
species to follow their suitable climates (12).
Lastly, effectiveness in controlling threats, such
as invasive species, encroachment, poaching, and
other impacts on protected lands, also critically
depends on the surrounding matrix (10). Thus, to
stem the tide of biodiversity loss, we must expand
beyond protected areas, using working lands con-
servation both to buffer and to reduce the threats
that cross park boundaries and to create acces-
sory habitats for both movement and persistence.
Working lands conservation is a key linchpin
for combatting the triple Anthropocene chal-
lenge of biodiversity loss, climate change, and
unsustainable land use. A large-scale example is
the Mesoamerican Biological Corridor project,
which 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 2of9
Fig. 1. Rebuilding connectivity in the matrix by using silvopasture. Photo of Finca La Luisa
showing several types of silvopastoral systems, including regenerating secondary tropical dry forest
trees with grass understory (yellow) and rows of planted Eucalyptus trees interspersed with
nitrogen-fixing Leucaena leucocephala fodder shrubs and forage grasses (blue). These systems were
established on former monoculture agricultural lands to restore compacted, degraded soils; the
red area shows early stages of tropical dry forest regeneration prior to grass seeding for silvopasture.
Silvopastures produce more cattle sustainably on less land, buffer ranchers from economic losses
due to climate extremes, and create landscape connectivity to other forest fragments (orange) in the
Cesar river valley, Colombia.
RESEARCH |REVIEW
PHOTO: J. J. LOPER A/CIPAV; ADAPTED BY N IRJA DESAI/SCIENC E
on October 18, 2018 http://science.sciencemag.org/Downloaded from
more than 650protected areas in the region (43).
A concurrent goal is to use sustainable agriculture
and forestry techniques to promote livelihoods
and enhance resilience to climate change (36).
Protected areas are vital in this region because
many species are restricted to forest; however, most
reserves are small and isolated. In combination
with steep elevational and latitudinal gradients
in the region, this isolation makes species in-
habiting reserves particularly vulnerable to climate
change. The Mesoamerican Biological Corridor
project recognizes the role that working lands can
play to restore critical connectivity by increasing
tree diversity and cover through live fences, agro-
forestry, silvopasture, forest fallows, home gardens,
and protection or restoration of riparian forests
and forest fragments (43). These forest elements,
which include both ribbonlike and patch struc-
tures, support a large number of neotropical
birds, insects, mammals, and plants (17,44); en-
hance the movement of birds and bats across the
landscape (45–47); and thus contribute to con-
servation, even of vulnerable wildlife (17,47,48).
Forest elements also promote sustainable land
use and contribute to local livelihoods by sup-
porting ecosystem services. For example, evi-
dence suggests that an economically devastating
invasive pest, the coffee berry borer, is reduced by
the integration of forest elements within coffee
landscapes, which both limits the borer’sability
to colonize new coffee fields (49) and promotes
bird species that prey on the borer (50). Reduced
economic losses due to pest control from birds are
similar in magnitude to average per capita income
in the region and are strongly related to forest
cover (50). Adopting sustainable agricultural
techniques and enhancing tree cover simulta-
neously creates more flexible and resilient pro-
ductionsystems that allow farmers and ranchers
to adapt to extreme conditions prompted by
climate change (33,51). Although some crit ics
decry the effectiveness of the Mesoamerican
Biological Corridor project, it may be too early to
judge. Quite a few integrated landscape initiatives
areconcentratedintheregion,inassociationwith
biological corridors (36). However, many began
relatively recently, and we know from the few
scientific studies that exist that developing an
effective multistakeholder participatory process
takes substantial time (36,43,52). In one case
that is more advanced (the San Juan–La Selva
Biological Corridor in Costa Rica), some success
has been achieved in arresting deforestation and
encouraging tree planting, forest regeneration,
and connectivity through a government-run pay-
ments for ecosystem services program, as well as
other grassroots initiatives (43,53).
Mechanisms for promoting working
lands conservation
The challenge of shifting from managing work-
ing lands solely for profit to conservation of
working lands is not insignificant, but there are
clear paths toward larger-scale integration of
this approach. These strategies include various
regulatory, voluntary, incentive, market-based,
or governance instruments (table S1), which vary
in their applicability to private, communal, or
state-owned lands and the extent to which they
support biodiversity conservation versus liveli-
hoods or economies (Fig. 3A). Each approach has
challenges, especially around reconciling conser-
vation and socioeconomic objectives (table S1)
(42,54). Collectively, problems associated with
regulatory and incentive programs can include
inter alia lack of permanence or compliance, com-
plex implementation, unintended economic con-
sequences, low adoption rates, high monitoring
costs, and little evaluationofeffectivenessagainst
goals (table S1).
Further, there is often the risk that the bio-
diversity conserved through these actions is not
equivalent to that which was lost because of eco-
nomically driven land conversion. Instruments
for private lands may result in piecemeal land
management actions that have little positive ef-
fect on biodiversity at the landscape scale; promis-
ing public-private initiatives to overcome this
defect include corridor planning (43,55) (Box 2
and Fig. 4) and landscape-level mitigation (table
S1). For example, landowners required to set aside
forest on their properties under Brazil’s forest
code may develop these lands in exchange for mit-
igating lands elsewhere within the same biome
that provide greater conservation value (56). Man-
aging the matrix to promote biodiversity could
also exacerbate human-wildlife conflict; how-
ever, the recovery of carnivore populations within
human-dominated areas in Europe provides a
hopeful and inspiring example for how landscapes
can be shared between wildlife and people (14)
(Box 3). These instruments can exacerbate the
unequal distribution of benefits and costs within
and across communities (table S1). For example,
trading development rights on forestlands in ex-
change for permitting high-density urban devel-
opment elsewhere can provide open spaces for
working lands conservation. However, such trades
could exacerbate the lack of access to open space
already experienced by low-income urban house-
holds. Thus, the effects of conservation measures
on social equity and environmental justice should
also be considered (57). A final concern is that
there is often a trade-off between the rigor of
environmental standards or restrictions enforced
and the likelihood of adoption (table S1); incen-
tive schemes that are flexible, provide obvious
Kremen et al., Science 362, eaau6020 (2018) 19 October 2018 3of9
A
B
C
Crops
Healthy
soils
Freshwater
Carbon
sequestration
Pest control
services
Pollination
services
Biodiversity
Connectivity
Forest
products Livestock
production
Biodiversity Crops
Healthy
soils
Freshwater
Carbon
sequestration
Pest control
services
Pollination
services
Connectivity
Forest
products Livestock
production
Monoculture row-crop Mixed cultivated, forest and range landscape
Rangelands
Diversied farm
Riparian forest
Fig. 2. Ecosystem ser vice trade-offs with land management. Radar diagrams display how
different land uses affect various ecosystem services and biodiversity. (A) Monoculture row cropping
contributes to food production at the expense of other ecosystem services and biodiversity. (B)Ina
working landscape managed for conservation, patch types differ in the services they provide, but each
patch type should display a relatively even array of services, minimizing trade-offs. (C) Across
patches, the services provided for the working landscape in (B) are multifunctional.
RESEARCH |REVIEW
ILLUSTRATION: N IRJA DESAI/SCIEN CE BASED ON C. KRE MEN AND A.M. MERENLE NDER; PHOTO: NATIONAL AERI AL IMAGERY PROGRA M, ADAPTED BY NIRJA DESAI/ SCIENCE
on October 18, 2018 http://science.sciencemag.org/Downloaded from
benefits, target likely adopters, fit the sociocul-
tural context, foster enabling market and reg-
ulatory environments, and provide technical
assistance may boost adoption (58). For example,
payments for conserving or restoring forests in
CostaRicaarebasedonarea,whereastransaction
costs are the same regardless of size, disincen-
tivizing smaller landowners from participating
in the payments for ecosystem services scheme.
Encouraging smallholders to participate would
require adjusting the costs of participation so
that these landowners could also realize net gains
(53). Although numerous changes are required,
careful attention to the construction of these
programs could increase their success.
Further, several current trends favor working
lands conservation approaches. First, new policy
instruments [such as REDD+ (Reducing Emis-
sions from Deforestation and Forest Degrada-
tion)] operating across a range of scales, from
individual private landholdings to large-scale
community-based or government-funded initia-
tives, are being developed to incentivize conser-
vation on working lands. Second, the number and
variety of institutions involved in working lands
conservation are increasing, and such institu-
tions include both public-private partnerships
and nongovernmental conservation organiza-
tions that formerly focused primarily on pro-
tected areas (36,59,60). Third, these institutions
can take advantage of recent increases in both
public and private “investments for conserva-
tion”(investments designed to cogenerate finan-
cial returns and conservation benefits) (60). Such
investments include projects in sustainable food
and fiber production, water quality and quantity
projects, and outright habitat conservation (in
the latter, financial returns are based onchang-
ing land values or carbon stocks). Fourth, out-
side of these investments, an increasing number
of companies have committed to greening their
supply chains by reducing the environmental
impacts at the source, processing, delivery, and
end-of-life management of the product (61). Al-
though supply chain greening requires much
better monitoring, accountability, and inclusion
of biodiversity conservation as an explicit goal
(61,62), it could ultimately contribute to conser-
vation in working landscapes, particularly given
the vast economic power represented within cor-
porations (61). A final trend is the creation of
voluntary, community-driven programs (Box 2)
in which local communities participate in the
conservation of working landscapes to gain in-
creased access to information and expertise, build
interpersonal connections, and obtain both per-
sonal benefits and public recognition for practic-
ing sustainable methods (63).
We argue that this latter trend of community-
basedactionsandtheinnovations,networks,
and social movements that sometimes emerge
from them present the most exciting opportunity
to turn the tide against the triple Anthropocene
threat [see also (64)]. Communities seeking solu-
tions for socioecological resilience frequently rely
on working lands conservation approaches. For
example, Sustainable Solutions restores man-
grove forests in Sri Lanka and India through
youth-based community engagement to build
shoreline resilience to cyclones while enhancing
livelihoods from fisheries dependent on man-
grove ecosystems.
Further, local initiatives can link together to
form larger networks with the help of boundary
organizations to form social movements that can
advance environmental policies, improve sus-
tainable behaviors, and demand supply chain
Kremen et al., Science 362, eaau6020 (2018) 19 October 2018 4of9
Private
Public
Intensive agriculture Conservation easement Private
reserve
Management incentives
Certication schemes
Mitigation
Payment for ecosystem services
Community based natural resources management
Indigenous conserved areas
Protected areas (IUCN Cat V,VI)
Government lands
Government
plantation
Protected areas
(IUCN Cat I–IV)
Development
Cultivated lands
Local diversication and landscape-scale heterogeneity
Single-species
patures
Rangelands (<10% 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
crop/livestock
Complex
rotation/
intercrop
Management
intensive
grazing
Restored
range
Nomadic
pastoralism
Agroforestry
system
Home garden
Native/non-native
multi-age forests
Restored
forest
Native forest
with EBM
Chemical intensication
Fig. 3. Approaches for conservation of working lands occupy the space (yellow) between highly
developed (brown) and highly conserved (green) land uses. (A) An array of tools are available
for working lands conservation, for private, communal, or public lands (see table S2 for more detail and
examples). 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 of
diversification (left to right) or chemical intensification (right to left). Cultivated lands include all
planted 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 the
mid-1980s to conserve biodiversity and sustain agriculture in Australia, resulting in more than
5000 Landcare and Coastcare groups. More than 20 countries have since adopted the model.
In Australia, this model combines substantial government investment with landowner and
community engagement. For example, Landcare groups across eastern Australia contribute to
the delivery of the Great Eastern Ranges (GER) Initiative (105), alongside public land management
authorities, conservation organizations, research institutions, and traditional owners groups.
The GER is one of Australia’s largest public-private partnerships to conserve biodiversity in the
face of climate change (Fig. 4) as part of Australia’s National Wildlife Corridors Plan. Landcare
groups along the corridor undertake restoration and management activities, along with community
building and engagement. In the Queanbeyan Landcare group, 25 landholders signed up to
increase the foraging habitat for the glossy black cockatoo (Calyptorhynchus lathami) through
the restoration of 10,000 she-oaks (Allocasuarina sp.) in production lands along three river
catchments.The social networks and learning spaces created are promising ways of encouraging
conservation commitment among land managers. However, far more landowners must become
engaged to restore connectivity at the scale desired.
RESEARCH |REVIEW
ILLUSTRATION: N IRJA DESAI/SCIEN CE BASED ON C. KRE MEN AND A.M MERENLEN DER
on October 18, 2018 http://science.sciencemag.org/Downloaded from
accountability (64). For example, the withdrawal
of the United States from the Paris Agreement
at the 21st Conference of Parties (COP21) and
delays in regulation of emissions by other nations
galvanized a series of on-the-ground climate
actions from civil society, businesses, nonpro-
fits, and subnational government. The Global
Action Climate Summit of 2018 instigated by
California governor Jerry Brown illustrates a
new stage of this growing social movement.
Its Land and Ocean Stewardship “30 × 30”chal-
lenge brings together more than 100 organ-
izations focused on managing forests, farmlands,
and oceans to provide 30% of the climate change
solution by 2030, rather than waiting on agree-
ments among nation states that continue to
fall short of the necessary carbon reduction
targets. The land management techniques
being developed locally to mitigate and adapt
to climate change are generally consistent with
the conservation of working lands approach
[e.g., (65)].
The benefits of local land conservation can
also be scaledup and made more effectiveif they
are carried out within a landscape or regional
conservation program organized by a state or
nonprofit agency (58). Innovative social and
institutional arrangements for working lands
conservation may emerge, such as The Nature
Conservancy’s BirdReturns program in California.
Through a reverse auction, the program finds and
pays farmers willing to alter water management
to create “pop-up”wetlands to provide habitats
for shorebirds during their northward migra-
tion, selecting sites that optimize the conserva-
tion benefits relative to payments (15).
Management techniques for conserving
working lands
Cultivated lands
Cultivated lands make up 12% of the terrestrial
ice-free surface (66) and comprise row and forage
crops, seeded pastures, vineyards and orchards,
mixed crop and livestock systems, and tree crops
and plantations (Fig. 3B). Cultivated lands are
often highly simplified ecologically; thus, they
rely extensively on chemical fertilizers and pes-
ticides to replace ecosystem services formerly
generated within or around agroecosystems (31),
often creating negative consequences for the
environment 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 by
fostering biophysical conditions and ecolog-
ical interactions favorable to crop production
(31,67,68), producing a more balanced (sustain-
able) distribution of ecosystem services (Fig. 2B).
Evidence also suggests that they minimize many
of the negative environmental consequences as-
sociated with simplified farming (31) (Fig. 5). Fur-
ther, these techniques can maintain crop yields
and profitability; create new market opportunities;
enhance food security, nutrition, and livelihoods;
and contribute substantially to the global food
supply, particularly under a changing climate
(table S2). Be cause they re ly on relatively low-
cost, low-technology, knowledge-based methods
(69), agroecological diversification techniques can
be made accessible to the majority of farmers.
[Small-scale farms with <5 ha make up 94% of
farms worldwide (40) and produce more than
halfofworldfoodcrops(70).] These farming
methods use open-pollinated seed varieties that
can be saved and cultivars that are locally adapted;
thus, they are less dependent on purchased seeds
and other inputs that can lead to poverty traps
(71). Multiple grassroots organizations and social
movements support learning, sharing, and adapta-
tion of agroecological knowledge and seeds
through farmer-to-farmer networks under par-
ticipatory governance (64). Diversified, agroeco-
logical practices are therefore farming methods
that are highly compatible with working lands
conservation, although potentially more ap-
pli cable t o certain farming systems. Large-scale
Kremen et al., Science 362, eaau6020 (2018) 19 October 2018 5of9
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. 4. The GER Corridor Initiative, Australia. The light green outline represents the plan to protect
and restore more than 3,600 km
2
as a climate corridor. The numbered, dark green shapes denote
regional alliances of conservation and natural resource management organizations, including
Landcare communities (Box 2). In the photo, members of the Molonglo Catchment Group Landcare
community conduct restoration.
Box 3. Carnivore conservation in shared landscapes.
Maintaining populations of large carnivores ranks among the greatest of conservation
challenges. These area-demanding species require larger territories than most protected areas
possess, potentially necessitating costly translocations to ensure gene flow and maintain
populations. Further, these species conflict with people in surrounding matrices through
predation on livestock or, occasionally, maiming or killing of humans. Nonetheless, in Europe,
most large carnivore populations are stable or expanding. One-third of the area of mainland
Europe hosts at least one permanent population of its four large carnivore species, persisting
alongside moderate human densities and largely outside of protected areas.The success of
carnivore conservation in Europe is attributed to well-enforced, coordinated legislative protection,
improvements in habitat and ungulate prey base, and rural depopulation. Importantly, ranchers
have found ways to live with carnivores by using carnivore-proofed electric fence s and re-
invigorating traditional livestock-guarding practices using shepherds and dogs (14). Similarly, in a
cultivated region in India, large carnivore species (the leopard and striped hyena) persist with
few conflicts despite high human densities (300 people/km
2
) and the lack of wild prey (106),
suggesting the potential that exists for carnivore conservation in shared landscapes.
RESEARCH |REVIEW
MAPS: GRE AT EASTERN RA NGES; PHOTO: ANDREW CAMP BELL; ADAPTED BY NIR JA DESAI/SCIENCE
on October 18, 2018 http://science.sciencemag.org/Downloaded from
commercial farmers that have invested heavily in
the machinery associated with chemically intensive
agriculture may not readily switch to agroeco-
logical techniques (68,72); however, the use of
some agroecological techniques can be compa-
tible with existing infrastructure and can lead
to reduced agrochemical use at similar or even
enhanced profits [e.g.,( 73)].
A concern is that the use of “wildlife-friendly”
agroecological practices will require more land
to be farmed to produce the same amount of
food, promoting deforestation and harming bio-
diversity (74). However, a number of diversified,
agroecological farming methods maintain or in-
crease yields (table S2) (32,50,73,75–78). For
example, techniques such as intercropping, cover
cropping, and crop rotation may promote crop
yields through a variety of ecological mechanisms
(23), including complementarity of water and
nutrient use (e.g., different crops access differ-
ent soil layers for water and nutrient uptake),
facilitation of nutrient uptake [e.g., intercropped
faba bean acidifies the soil, mobilizing phospho-
rus that is taken up by rice (79)], reduction of
pests and diseases [e.g., pests and diseases spread
more slowly in spatially or temporally heteroge-
neous crop systems, and such systems also sup-
port predator populations that keep pests in
check (80,81)], and enhancement of soil biota
and fertility (82). By improving soil structure and
stability, which then enhances water infiltration
and retention, these techniques also stabilize
yields against annual environmental fluctua-
tions and more catastrophic disturbances such
as droughts and hurricanes (32,33).
Beyond providing resources and habitats for
agrobiodiversity, spe cific techniques such as agro-
forestry and the use of silvopasture, hedgerows,
flower strips, live fences, and riparian buffers
mayalsoenhancetheconnectivityoflandscapes
and promote the dispersal of various wildlife
species (16,47,83). Although these structural
features are known to increase the occurrence of
a wide variety of organisms within agricultural
landscapes (43,84), how they affect the dispersal
potential of organisms within diversified agri-
cultural lands is poorly understood. Nonetheless,
ambitious, large-scale connectivity projects, such
as the Mesoamerican Biological Corridor project
(43), the silvopastoral and rotational grazing proj-
ect in the Santa Catarina Atlantic Forest (55),
variouslinkagesinAustralia(Box2),andtheres-
toration of the migratory pathway of the mon-
arch butterfly (Danaus plexippus) in the U.S.
midwestern states (85),areunderwayforagricul-
turallands.Inthelattercase,althoughadaunt-
ing amount of restoration would be required to
support the butterfly, it could simultaneously en-
hance soybean pollination, improve water quality,
protect 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 and make transformation
to diversified, agroecological systems difficult
(68,72), reasons for optimism exist. Global grass-
roots movements such as La Via Campesina have
provided technical, social, and material support
to farmers for the spread of agroecology, con-
fronted industrial agribusiness, and fought to
influence national and global policies(64). Alter-
native agrifood systems and local and regional
initiatives that provide support for diversified,
agroecological systems are emerging (64,69). In-
ternational initiatives supporting agroecology
include the United Nations Right to Food program,
which embraces it as a key element for enhancing
food security globally (88), and programs of the
Food and Agriculture Organization, which has
held global and regional conferences on agro-
ecology and included it in Farmer Field Schools
since 2014 (68).
Rangelands and forests
Forests in the boreal, temperate, and tropical
regionsmakeup~30%ofEarth’sarea(89),
whereas rangelands, which are defined as having
<10% tree cover and include grasslands, desert
shrublands, savanna woodlands, alpine meadows,
and areas of tundra grasses and shrubs, constitute
~44% (90). Grazed by wild and domestic animals,
they vary greatly in productivity. Both natural
forests and rangelands have been lost or degraded
over the past several hundred years by the in-
creased extent and intensity of human use, in-
cluding timber harvest, grazing, and conversion
to agriculture. Forests continue to be lost and
degraded at an alarming rate (62), although for-
est regrowth due to rural depopulation is also
occurring in some areas (20). A recent global
analysis of so urces of tree cover losses showed
that industrial agriculture for commodity crops
is responsible for the permanent conversion of
5 million ha of forest per year (27% of losses, con-
centrated primarily in portions of Latin America
and Southeast Asia), whereas shifting agriculture
(primarily in Africa) and forestry (primarily in
North America and Europe) cause forest distur-
bance or degradation over an equivalent land
Kremen et al., Science 362, eaau6020 (2018) 19 October 2018 6of9
Monoculture crop: Adding prairie strips (10%):
8 inches/acre runo
4 tons/acre sediment lost
7 lbs/acre phosphorus lost
35 lbs/acre nitrogen lost
42% less runo
95% less soil export
89% less phosphorus export
84% less nitrogen export
Fig. 5. Diversification practices can increase biodiversity. The integration of prairie strips into a
corn-soy rotation exemplifies how diversification within working lands can substantially increase plant,
pollinator, and bird species richness and abundance by two- to fourfold (as indicated by colors and
numbers of icons, respectively) while minimizing externalities and enhancing other ecosystem
services, such as pollination for the soy crop (table S2) (86).
RESEARCH |REVIEW
ILLUSTRATION: N IRJA DESAI/SCIEN CE BASED ON STRIP S RESEARCH TEAM AND LEO POLD CENTER FOR SUSTAINAB LE AGRICULTURE
on October 18, 2018 http://science.sciencemag.org/Downloaded from
area, followed by regrowth (62). It is critical,
therefore, to cease permanent conversion of
forests for commodity cropping and to apply
restorative management approaches in working
forests and rangelands.
Since1990,manynationshavecreateden-
abling policies and legislation for sustainable
forest management (89). Of the 54% of global
forests considered “permanent”(that is, expected
to retain forest cover in the long term), 99% of
these 2.17 billion ha are covered by such policies,
a necessary but not sufficient condition for sus-
tainable management. Indicators of sustainable
management also show positive temporal trends,
but over smaller areas. For example, forest cer-
tification (table S1) covered 430 million ha by
2014 (89), but largely within boreal and temper-
ate regions, where land-clearing rates are less
acute than those in the tropics.
An array of restorative fore st and rangeland
management options exist that are compati-
ble with the conservation of working lands (Fig.
3B and table S2). For forests, the adoption of
ecosystem-based managementapproaches has
led to the integration of a greater variety of tree
species and age and size classes, including old
growth 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 maintains
similarities between natural and managed forests,
contrasting with even-aged management from
clear-cutting. Evidence from silvicultural trials and
natural forests suggests that greater tree diversity
also enhances wood yield quantity and stability
(23). In keeping with the ecosystem stewardship
concept (27), ecosystem-based management also
emphasizes collaborative decentralized control
and adaptive management, as well as landscape
planning and the designation of corridors to pro-
mote wildlife (92). However, stakeholders may
reject harvesting practices that negatively af-
fec t financial returns in the short term. Environ-
mental outcomes suffered when stakeholders
had stronger oversight of the process than a
regulatory authority with political backing (93),
support ing the need for public-private part-
nerships to achieve biodiversity conservation
objectives.
In rangelands, compatible management prac-
tices are exemplified by the dehesa and montado
traditional pastoral systems in oak savannas of
Spain and Portugal, respectively. The oak trees
(Quercus rotundifolia and Q. suber)arepruned
to increase the production of acorns to feed to
pigs and other livestock grown for high-value
meat 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 Europe.
However, grazing, browsing, and trampling can
limit oak regeneration; thus, pasture areas need
periodic temporary protection from livestock to
promote oak recruitment and sustainable use
(95). In Colombia, many ranchers are restoring
degraded agricultural lands by using various
silvopastoral techniques, which also enhance
connectivity in these landscapes (Fig. 1).
Freshwater ecosystems
Maintaining stream flows and hydrologic con-
nectivity is essential for conserving freshwater
biodiversity and ecosystems. Because of changes
in stream flows, estimates suggest that up to 75%
of freshwater fish species are headed for local
extinction by 2070 (96). Fresh water also limits
the production of many natural resources, and
its quantity and quality are in turn affected by
landscape management. Appropriate manage-
ment techniques can promote groundwater re-
charge and stream flow in working landscapes
(table S2) (31,86), of increasing importance
under drier futures with more extreme precip-
itation events (97). Flood plains and associated
riparian zones are particularly critical to conserve
in working landscapes, because they dispropor-
tionately support biodiversity and ecosystem
processes compared with other landscape ele-
ments (98). Riparian corridors also provide cooler
and moister microclimates than surrounding areas
and often span elevational and climatic gradients
that may permit species to follow their climate
envelopes (99).
Recommendations and
concluding thoughts
Managing the working lands matrix for bio-
diversity needsto become a mainstream compo-
nent of public and private conservation efforts,
complementing the more traditional (and essen-
tial) focus on increasing the extent and effective-
ness of protected areas (16). These restorative,
working lands conservation approaches (table
S2)shouldbeappliedtothelargelandareathat
is already used for farming, forestry, and ranch-
ing. At the same time, we critically need policies
to prevent further conversion and degradation of
wilderness and relatively intact ecosystems (62).
To scale up working lands conservation, in-
creased support is needed for the voluntary, policy,
and market instruments described in table S1.
However, further adaptation and learning is
needed to improve their efficacy, both at the
project level and through evidence-based synthe-
ses [e.g., (100], and to increase adoption rates
by considering an array of social factors (58).
Further, these measures must be complemented
by community-driven conservation initiatives,
which, by involving young and old in steward-
ship, communication, citizen science, and edu-
cation, can create a shared vision and innovative
practicesthatresultincollectiveimpact.Scien-
tists can support community-driven conserva-
tion and help advance environmental social
movements by engaging the public, listening to
alternative ways of knowing, and cocreating con-
servation, management, and policy alternatives.
Especially important is to create alliances with
existing community actions and social movements
that share common ground, such as climate or
local food movements.
Ultimately, our efforts to protect biodiversity
and sustain resources must be accompanied by
measures to reduce human population and con-
sumption while increasing equitable access to
resources to achieve sustainability. Opportunities
to stabilize population and consumption exist.
For example, through concerted government in-
vestment in voluntary family planning programs,
enormous progress in reducing total fertility
rates has been made even in poor countries [e.g.,
(101], leading to smaller families living better.
Globally, a large unmet need for family planning
still exists (101); further investment could help
stabilize the global population at 6 billion people
by 2100, instead of the 9 to 12 billion projected
without intervention (102,103). To reduce con-
sumption, critical targets include reducing food
waste and meat consumption (104) and seeking
efficiencies in energy and water use that can
accompany urbanization (102). Even with well-
structured policies, these changes toward lower
human population and consumption would take
time; thus, concerns exist that humanity will
destroy biodiversity and natural resources before
achieving a more sustainable human population
(102). Conservation in working landscapes can
help maintain all species, including people, as we
strive to achieve a planet where a smaller human
population lives better and more equitably with
and because of wild nature.
REFERENCES AND NOTES
1. J. L. Payne, A. M. Bush, N. A. Heim, M. L. Knope,
D. J. McCauley, Ecological selectivity of the emerging mass
extinction in the oceans. Science 353, 1284–1286 (2016).
doi: 10.1126/science.aaf2416; pmid: 27629258
2. G. Ceballos, P. R. Ehrlich, R. Dirzo, Biological annihilation via
the ongoing sixth mass extinction signaled by vertebrate
population losses and declines. Proc. Natl. Acad. Sci. U.S.A.
114, E6089–E6096 (2017). pmid: 28696295
3. G. Ceballos et al., Accelerated modern human-induced species
losses: Entering the sixth mass extinction. Sci. Adv. 1, e1400253
(2015). doi: 10.1126/sciadv.1400253;pmid:2 6601195
4. C. A. Hallmann et al., More than 75 percent decline over
27 years in total fly ing insect biomas s in protected area s.
PLOS ONE 12, e01858 09 (2017). doi: 1 0.1371/
journal.pone. 0185809;pmid:29045418
5. T. Newbold et al., Global effects of land use on local
terrestrial biodiversity. Nature 520,45–50 (2015).
doi: 10.1038/nature14324; 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 mountain
ranges. Proc. Natl. Acad. Sci. U.S.A. 115, 6004–6009 (2018).
doi: 10.1073/pnas.1720141115; pmid: 29784825
8. R. M. Pringle, Upgrading protected areas to conserve wild
biodiversity. Nature 546,91–99 (2017). doi: 10.1038/
nature22902; pmid: 28569807
9. J. M. Halley, N. Monokrousos, A. D. Mazaris, W. D. Newmark,
D. Vokou, Dynamics of extinction debt across five taxonomic
groups. Nat. Commun. 7, 12283 (2016). doi: 10.1038/
ncomms12283; pmid: 27452815
10. W. F. Laurance et al., Averting biodiversity collapse in tropical
forest protected areas. Nature 489, 290–294 (2012).
doi: 10.1038/nature11318;pmid: 22832582
11. M. C. Urban, Climate change. Accelerating extinction risk
from climate change. Science 348, 571–573 (2015).
doi: 10.1126/science.aaa4984; pmid: 25931559
12. S. R. Loarie et al., The velocity of climate change. Nature 462,
1052–1055 (2009). doi: 10.1038/nature08649;
pmid: 20033047
13. L. O. Frishkoff et al., Loss of avian phylogenetic
diversity in neotropical agricultural systems. Science
345, 1343–1346 (2014). doi: 10.1126/science.1254610;
pmid: 25214627
14. G. Chapron et al., Recovery of large carnivores in Europe’s
modern human-dominated landscapes. Science 346,1517–1519
(2014). doi: 10.1126/science.1257553; pmid: 25525247
Kremen et al., Science 362, eaau6020 (2018) 19 October 2018 7of9
RESEARCH |REVIEW
on October 18, 2018 http://science.sciencemag.org/Downloaded from
15. M. D. Reynolds et al., Dynamic conservation for migratory
species. Sci. Adv. 3, e1700707 (2017). doi: 10.1126/
sciadv.1700707; pmid: 28845449
16. C. Kremen, Reframing the land-sparing/land-sharing
debate for biodiversity conservation. Ann. N.Y. Acad. Sci.
1355, 52–76 (2015). doi: 10.1111/nyas.12845;
pmid: 26213864
17. C. D. Mendenhall, A. Shields-Estrada, A. J. Krishnaswami,
G. C. Daily, Quantifying and sustaining biodiversity in tropical
agricultural landscapes. Proc. Natl. Acad. Sci. U.S.A. 113,
14544–14551 (2016). doi: 10.1073/pnas.1604981113;
pmid: 27791070
18. C. D. Mendenhall, D. S. Karp, C. F. J. Meyer, E. A. Hadly,
G. C. Daily, Predicting biodiversity change and averting
collapse in agricultural landscapes. Nature 509, 213–217
(2014). doi: 10.1038/nature13139; pmid: 24739971
19. C. M. Kennedy, E. H. C. Grant, M. C. Neel, W. F. Fagan,
P. P. Marra, Landscape matrix mediates occupancy dynamics
of Neotropical avian insectivores. Ecol. Appl. 21, 1837–1850
(2011). doi: 10.1890/10-1044.1; pmid: 21830722
20. J. A. Foley et al., Global consequences of land use. Science
309, 570–574 (2005). doi: 10.1126/science.1111772;
pmid: 16040698
21. N. Ramankutty et al., Trends in global agricultural land use:
Implications for environmental health and food security.
Annu. Rev. Plant Biol. 69, 789–815 (2018). doi: 10.1146/
annurev-arplant-042817-040256; pmid: 29489395
22. F. Isbell et al., Linking the influence and dependence
of people on biodiversity across scales. Nature 546,65–72
(2017). doi: 10.1038/nature22899; pmid: 28569811
23. F. Isbell et al., Benefits of increasing plant diversity in
sustainable agroecosystems. J. Ecol. 105,871–879 (2017).
doi: 10.1111/1365-2745.12789
24. D. J. Eastburn, A. T. O’Geen, K. W. Tate, L. M. Roche, Multiple
ecosystem services in a working landscape. PLOS ONE 12,
e0166595 (2017). doi: 10.1371/journal.pone.0166595;
pmid: 28301475
25. D. A. Driscoll, S. C. Banks, P. S. Barton, D. B. Lindenmayer,
A. L. Smith, Conceptual domain of the matrix in fragmented
landscapes. Trends Ecol. Evol. 28, 605–613 (2013).
doi: 10.1016/j.tree.2013.06.010; pmid: 23883740
26. W. F. Fagan, E. E. Holmes, Quantifying the extinction vortex.
Ecol. Lett. 9,51–60 (2006). pmid: 16958868
27. F. S. Chapin 3rd et al., Ecosystem stewardship: Sustainability
strategies for a rapidly changing planet. Trends Ecol. Evol. 25,
241–249 (2010). doi: 10.1016/j.tree.2009.10.008;
pmid: 19923035
28. F. Isbell et al., High plant diversity is needed to maintain
ecosystem services. Nature 477, 199–202 (2011).
doi: 10.1038/nature10282; pmid: 21832994
29. M. G. E. Mitchell, E. M. Bennett, A. Gonzalez, Linking
landscape connectivity and ecosystem service provision:
Current knowledge and research gaps. Ecosystems 16,
894–908 (2013). doi: 10.1007/s10021-013-9647-2
30. M. Loreau, N. Mouquet, A. Gonzalez, Biodiversity as spatial
insurance in heterogeneous landscapes. Proc. Natl. Acad. Sci.
U.S.A. 100, 12765–12770 (2003). doi: 10.1073/
pnas.2235465100; pmid: 14569008
31. C. Kremen, A. Miles, Ecosystem services in biologically
diversified versus conventional farming systems: Benefits,
externalities, and trade-offs. Ecol. Soc. 17, 40 (2012).
doi: 10.5751/ES-05035-170440
32. A. C. M. Gaudin et al., Increasing crop diversity mitigates
weather variations and improves yield stability. PLOS ONE 10,
e0113261 (2015). doi: 10.1371/journal.pone.0113261;
pmid: 25658914
33. S. M. Philpott, B. B. Lin, S. Jha, S. J. Brines, A multi-scale
assessment of h urricane impact s on agricultur al landscapes
basedonlanduseandtopographicfeatures.Agric.
Ecosyst. Environ. 128,12–20 (2008). doi: 10.1016/
j.agee.2008.04.016
34. L. C. Ponisio et al., Diversification practices reduce organic to
conventional yield gap. Proc. R. Soc. London Ser. B 282,
20141396 (2015). doi: 10.1098/rspb.2014.1396;
pmid: 25621333
35. R. J. Hobbs et al., Managing the whole landscape: Historical,
hybrid, and novel ecosystems. Front. Ecol. Environ. 12,
557–564 (2014). doi: 10.1890/130300
36. N. Estrada-Carmona, A. K. Hart, F. A. J. DeClerck,
C. A. Harvey, J. C. Milder, Integrated landscape management
for agriculture, rural livelihoods, and ecosystem conservation:
An assessment of experience from Latin America and
the Caribbean. Landsc. Urban Plan. 129,1–1 1 (2014).
doi: 10.1016/j .landu rbplan. 2014.05. 001
37. H. Gu, S. M. Subramanian, Drivers of change in
socio-ecological production landscapes: Implications for
better management. Ecol. Soc. 19,41(2014).doi:10.5751/
ES-06283-190141
38. M. K. Anderson, Tending the Wild: Native American Knowledge
and the Management of California’s Natural Resources
(Univ. of California Press, 2005).
39. E. M. Bignal, D. I. McCracken, The nature conservation value
of European traditional farming systems. Environ. Rev. 8,
149–171 (2000). doi: 10.1139/a00-009
40. S. K. Lowder, J. Skoet, T. Raney, The number, size, and
distribution of farms, smallholder farms, and family farms
worldwide. World Dev. 87,16–29 (2016). doi: 10.1016/
j.worlddev.2015.10.041
41. C. L. Gray et al., Local biodiversity is higher inside than
outside terrestrial protected areas worldwide. Nat. Commun.
7, 12306 (2016). doi: 10.1038/ncomms12306;
pmid: 27465407
42. C. L. Shafer, Cautionary thoughts on IUCN protected area
management categories V-VI. Global Ecol. Conserv. 3,
331–348 (2015). doi: 10.1016/j.gecco.2014.12.007
43. F. A. J. De Clerck et al., Biodiversit y conservation in
human-modified landscapes of Mesoamerica: Past, present
and future. Biol. Conserv. 143, 2301–2313 (2010).
doi: 10.1016/j.biocon.2010.03.026
44. C. A. Ha rvey et al., Patterns of animal diversity
in different forms of tree cover in agricultural
landscapes. Ecol. Appl. 16,1986–1999 (2006).
doi: 10.1890/1051-0761(2006)016[1986:POADID]2.0.CO;2;
pmid: 17069389
45. Ç. H. Şekercioğlu et al., Tropical countryside riparian
corridors provide critical habitat and connectivity for
seed-dispersing forest birds in a fragmented landscape.
J. Ornithol. 156 (Suppl. 1), 343–353 (2015). doi: 10.1007/
s10336-015-1299-x
46. C. A. Harvey et al., Contribution of live fences to the
ecological integrity of agricultural landscapes.
Agric. Ecosyst. Environ. 111,200–230 (2005). doi: 10.1016/
j.agee.2005.06.011
47. A. Medina, C. A. Harvey, D. S. Merlo, S. Vílchez, B. Hernández,
Bat diversity and movement in an agricultural landscape in
Matiguás, Nicaragua. Biotropica 39, 120–128 (2007).
doi: 10.1111/j.1744-7429.2006.00240.x
48. K. Williams-Guillen, C. McCann, J. C. Martinez Sanchez,
F. Koontz, Resource availability and habitat use by mantled
howling monkeys in a Nicaraguan coffee plantation: Can
agroforests serve as core habitat for a forest mammal?
Anim. Conserv. 9, 331–338 (2006). doi: 10.1111/
j.1469-1795.2006.00042.x
49. J. Avelino, A. Romero-Gurdián, H. F. Cruz-Cuellar,
F. A. J. Declerck, Landscape context and scale differentially
impact coffee leaf rust, coffee berry borer, and coffee
root-knot nematodes. Ecol. Appl. 22, 584–596 (2012).
doi: 10.1890/11-0869.1; pmid: 22611856
50. D. S. Karp et al., Forest bolsters bird abundance, pest control
and coffee yield. Ecol. Lett. 16, 1339–1347 (2013).
doi: 10.1111/ele.12173; pmid: 23981013
51. C. A. Harvey et al., Climate-smart landscapes: Opportunities
and challenges for integrating adaptation and mitigation in
tropical agriculture. Conserv. Lett. 7,77–90 (2014).
doi: 10.1111/conl.12066
52. A. T. H. Keeley et al., Making habitat connectivity a
reality. Conserv. Biol. (2018). doi: 10.1111/cobi.13158
pmid: 29920775
53. W. C. Morse et al., Consequences of environmental service
payments for forest retention and recruitment in a Costa Rican
biological corridor. Ecol. Soc. 14, 23 (2009). doi: 10.5751/
ES-02688-140123
54. J. Owley, D. Takacs, “Flexible Conservation in Uncertain
Times,”in Contemporary Issues in Climate Change Law and
Policy: Essays Inspired by the IPCC (UC Hastings Research
Paper 180, Univ. at Buffalo School of Law, 2016),
pp. 65–102; https://papers.ssrn.com/sol3/papers.cfm?
abstract_id=2700658.
55. J. Alvez, A. L. Schmitt Filho, J. Farley, G. Alarcon, A. C. Fantini,
The potential fo r agroecosystem s to restore ecol ogical
corridors and sustain farmer livelihoods: Evidence from
Brazil. Ecol. Restor. 30,288–290 (2012). doi: 10.3368/
er.30.4.288
56. C. M. Kennedy et al., Bigger is better: Improved nature
conservation and economic returns from landscape-level
mitigation. Sci. Adv. 2, e1501021 (2016). doi: 10.1126/
sciadv.1501021; pmid: 27419225
57. I. Scoones, P. Newell, M. Leach, in The Politics of Green
Transformations, I. Scoones, M. Leach, P. Newell, Eds.
(Earthscan from Routledge, 2015), pp. 1–24.
58. M. B. Mascia, M. Mills, When conservation goes viral:
The diffusion of innovative biodiversity conservation
policies and practices. Conserv. Lett. 11, e12442 (2018).
doi: 10.1111/conl.12442
59. D. F. Doak, V. J. Bakker, B. E. Goldstein, B. Hale, What
is the future of conservation? Trends Ecol. Evol. 29,77–81
(2014). doi: 10.1016/j.tree.2013.10.013; pmid: 24332874
60. K. Hamrick, “State of private investment in conservation
2016: A landscape assessment of an emerging market”
(Forest Trends, 2016).
61. R. Chaplin-Kramer et al., Ecosystem service information to
benefit sustainability standards for commodity supply chains.
Ann. N.Y. Acad. Sci. 1355,77–97 (2015). doi: 10.1111/
nyas.12961; pmid: 26555859
62. P. G. Curtis, C. M. Slay, N. L. Harris, A. Tyukavina,
M. C. Hansen, Classifying drivers of global forest loss.
Science 361, 1108–1111 (2018). doi: 10.1126/science.
aau3445; pmid: 30213911
63. D. Pannell et al., Understanding and promoting adoption of
conservation practices by rural landholders. Aust. J. Exp.
Agric. 46, 1407–1424 (2006). doi: 10.1071/EA05037
64. M. Leach, I. Scoones, in The Politics of Green
Transformations, I. Scoones, P. Newell, M. Leach, Eds.
(Earthscan from Routledge, 2015), pp. 119–133.
65. D. R. Cameron, D. C. Marvin, J. M. Remucal, M. C. Passero,
Ecosystem management and land conservation can
substantially contribute to California’s climate mitigation
goals. Proc. Natl. Acad. Sci. U.S.A. 114, 12833–12838 (2017).
doi: 10.1073/pnas.1707811114; pmid: 29133408
66. N. Ramankutty, A. T. Evan, C. Monfreda, J. A. Foley, Farming
the planet: 1. Geographic distribution of global agricultural
lands in the year 2000. Global Biogeochem. Cycles 22,
GB1003 (2008). doi: 10.1029/2007GB002952
67. M. A. Altieri, The ecological role of biodiversity in
agroecosystems. Agric. Ecosyst. Environ. 74,19–31 (1999).
doi: 10.1016/S0167-8809(99)00028-6
68. International Panel of Experts on Sustainable Food Systems
(IPES-Food), “From uniformity to diversity: A paradigm
shift from industrial agriculture to diversified agroecological
systems”(IPES-Food, 2016); http://www.ipes-food.org/
images/Reports/UniformityToDiversity_FullReport.pdf.
69. C. Kremen, A. Iles, C. M. Bacon, Diversified farming systems:
An agro-ecological, systems-based alternative to modern
industrial agriculture. Ecol. Soc. 17, 44 (2012). doi: 10.5751/
ES-05103-170444
70. V. Ricciardi, N. Ramankutty, Z. Mehrabi, L. Jarvis,
B. Chookolingo, How much of the world’s food do
smallholders produce? Global Food Sec. 17,64–72 (2018).
doi: 10.1016/j.gfs.2018.05.002
71. M. Montenegro, Banking on wild relatives to feed the world.
Gastron. J. Crit. Food Stud. 16,1–8 (2016). doi: 10.1525/
gfc.2016.16.1.1
72. A. Iles, R. Marsh, Nurturing diversified farming systems
in industrialized countries: How public policy can
contribute. Ecol. Soc. 17, 42 (2012). doi: 10.5751/
ES-05041-170442
73. A. S. Davis, J. D. Hill, C. A. Chase, A. M. Johanns,
M. Lie bman, Increasing cropping system diversity balances
productivity, profitability and environmental health. PLOS
ONE 7, e47149 (2012). doi: 10.1371/journal.pone.0047149;
pmid: 23071739
74. B. Phalan, M. Onial, A. Balmford, R. E. Green, Reconciling food
production and biodiversity conservation: Land sharing and
land sparing compared. Science 333,1289–1291 (2011).
doi: 10.1126/science.1208742;pmid:21885781
75. K. Garbach et al., Examining multi-functionality for crop yield
and ecosystem services in five systems of agroecological
intensification. Int. J. Agric. Sustain. 15,11–28 (2017).
doi: 10.1080/14735903.2016.1174810
76. J.Pretty,Z.P.Bharucha,Sustainableintensificationin
agricultural systems. Ann. Bot. 114, 1571–1596 (2014).
doi: 10.1093/aob/mcu205; pmid: 25351192
77. G. M. Gurr et al., Multi-country evidence that crop diversification
promotes ecologi cal intensifica tion of agricult ure.
Nat. Plants 2, 16014 (2016). doi: 10.1038/nplants.20 16.14;
pmid: 27249349
78. R. F. Pywell et al., Wildlife-friendly farming increases crop
yield: Evidence for ecological intensification. Proc. R. Soc.
London Ser. B 282, 20151740 (2015). doi: 10.1098/
rspb.2015.1740; pmid: 26423846
Kremen et al., Science 362, eaau6020 (2018) 19 October 2018 8of9
RESEARCH |REVIEW
on October 18, 2018 http://science.sciencemag.org/Downloaded from
79. L. Li et al., Diversity enhances agricultural productivity via
rhizosphere phosphorus facilitation on phosphorus-deficient
soils. Proc. Natl. Acad. Sci. U.S.A. 104, 11192–11196 (2007).
doi: 10.1073/pnas.0704591104; pmid: 17592130
80. D. K. Letourneau et al., Does plant diversity benefit
agroecosystems? A synthetic review. Ecol. Appl. 21,9–21
(2011). doi: 10.1890/09-2026.1; pmid: 21516884
81. Y. Zhu et al., Genetic diversity and disease control in rice.
Nature 406, 718–722 (2000). doi: 10.1038/35021046;
pmid: 10963595
82. S. F. Bender, C. Wagg, M. G. A. van der Heijden, An
underground revolution: Biodiversity and soil ecological
engineering for agricultural sustainability. Trends Ecol. Evol. 31,
440–452 (2016). doi: 10.1016/ j.tree.2016.02 .016;pmid:
26993667
83. I. Perfecto, J. Vandermeer, A. Wright, Nature’sMatrix:Linking
Agriculture, Conservation and Food Sovereignty (Earthscan, 2009).
84. J. A. Hilty, A. M. Merenlender, Use of riparian corridors and
vineyards by mammalian predators in northern California.
Conserv. Biol. 18, 126–135 (2004). doi: 10.1111/j.1523-
1739.2004.00225.x
85. W. E. Thogmartin et al., Restoring monarch butterfly habitat
in the Midwestern US: ‘All hands on deck.’Environ. Res. Lett.
12, 074005 (2017). doi: 10.1088/1748-9326/aa7637
86. L. A. Sc hulte et al., Prairie strips improve biodiversity
and the delivery of multiple ecosystem services from
corn-soybean croplands. Proc. Natl. Acad. Sci. U.S.A. 114,
11247–11252 (2017). doi: 10.1073/pnas.1620229114;
pmid: 28973922
87. E. Brandes et al., Subfield profitability analysis reveals an
economic case for cropland diversification. Environ. Res. Lett.
11, 014009 (2016). doi: 10.1088/1748-9326/11/1/014009
88. O. De Schutter, “Agroecology and the right to food,”report
presented at the 16th Session of the United Nations
Human Rights Council [A/HRC/16/49], Geneva, Switzerland,
8 March 2011.
89. K. G. MacDicken et al., Global progress toward sustainable
forest management. For. Ecol. Manage. 352,47–56 (2015).
doi: 10.1016/j.foreco.2015.02.005
90. H. G. Lund, Accounting for the world’s rangelands.
Soc. Range Manage. 29,3–10 (2007).
91. S. Gauthier, M.-A. Vaillancourt, D. Kneeshaw, P. Drapeau,
L. De Grandpré, Y. Claveau, D. Paré, in Ecosystem Management
in the Boreal Forest, S. Gauthier, M.-A. Vaillancourt, A. Leduc,
L. De Grandpré, D. Kneeshaw, H. Morin, P. Drapeau,
Y. Bergeron, Eds. (Univ. du Québec, 2009), pp. 13–38.
92. M. Mangel et al., Principles for the conservation of wild living
resources. Ecol. Appl.6,338–362 (1996). doi: 10.2307/2269369
93. J. A. Layzer, Natural Experiments: Ecosystem-Based
Management and the Environment (MIT, 2008).
94. P. Campos, L. Huntsinger, J. L. Oviedo, P. F. Starrs, M. Diaz,
R. B. Standiford, G. Montero, Eds., Mediterranean Oak
Woodland Working Landscapes:Dehesas of Spainand Ranchlands
of California (Springer Science+Business Media, 2013).
95. J. A. Ramírez, M. Díaz, The role of temporal shrub
encroachment for the maintenance of Spanish holm oak
Quercus ilex dehesas. For. Ecol. Manage. 255, 1976–1983
(2008). doi: 10.1016/j.foreco.2007.12.019
96. M. A. Xenopoulos et al., Scenarios of freshwater fish extinctions
from climate change and water withdrawal. Global Change Biol.
11,1557–1564 (2005). doi:10.1111/j.1365-2486.2005.001008.x
97. S. D. Polade, A. Gershunov, D. R. Cayan, M. D. Dettinger,
D. W. Pierce, Precipitation in a warming world: Assessing
projected hydro-climate changes in California and other
Mediterranean climate regions. Sci. Rep. 7,10783(2017).
doi: 10.1038/s41598-017-11285-y;pmid:28883636
98. F. R. Hauer et al., Gravel-bed river floodplains are the
ecological nexus of glaciated mountain landscapes. Sci. Adv.
2, e1600026 (2016). doi: 10.1126/sciadv.1600026;
pmid: 27386570
99. A. T. Keeley et al., New concepts, models, and assessments
of climate-wise connectivity. Environ. Res. Lett. 13, 073002
(2018). doi: 10.1088/1748-9326/aacb85
100. W. Sutherland, L. V. Dicks, N. Ockendon, R. Smith, Eds., What
Works in Conservation (Open Book, ed. 2, 2017), vol. 2.
101. J. Bongaarts, S. W. Sinding, A response to critics of family
planning programs. Int. Perspect. Sex. Reprod. Health 35,
39–44 (2009). doi: 10.1363/3503909; pmid: 19465347
102. E. W. Sanderson, J. Walston, J. G. Robinson, From bottleneck
to breakthrough: Urbanization and the future of biodiversity
conservation. B ioscience 68,412–426 (2018). doi: 10.1093/
biosci/biy039;pmid:29867252
103. C. J. Bradshaw, B. W. Brook, Human population reduction is
not a quick fix for environmental problems. Proc. Natl.
Acad. Sci. U.S.A. 111, 16610–16615 (2014). doi: 10.1073/
pnas.1410465111; pmid: 25349398
104. J. Poore , T. Nemec ek, Reduc ing food ’s environmental
impacts through producers and consumers. Science
360,987–992 (2018). doi: 10.1126/science.aaq0216;
pmid: 29853680
105. Great E astern Ra nges, www.ger.org.au.
106. V. Athreya, M. Odden, J. D. C. Linnell, J. Krishnaswamy,
U. Karanth, Big cats in our backyards: Persistence of large
carnivores in a human dominated landscape in India.
PLOS ONE 8, e57872 (2013). doi:10.1371/journal.pone.0057872;
pmid: 23483933
ACKNO WLED GME NTS
We appreciate the constructive input of D. Ackerly, B. Brunner,
A. Campbell, F. DeClerck, and A. Knight. Competing interests: The
authors declare no competing interests.
SUPPLEMENTARY MATERIALS
www.sciencemag.org/content/362/6412/eaau6020/suppl/DC1
Tables S1 and S2
References (107–158)
10.1126/science.aau6020
Kremen et al., Science 362, eaau6020 (2018) 19 October 2018 9of9
RESEARCH |REVIEW
on October 18, 2018 http://science.sciencemag.org/Downloaded from
Landscapes that work for biodiversity and people
C. Kremen and A. M. Merenlender
DOI: 10.1126/science.aau6020
(6412), eaau6020.362Science
, this issue p. eaau6020Science
just for ecosystem services but also for maintenance and persistence of nonhuman species.
most human-modified lands as ''working landscapes.'' These can provide for human needs and maintain biodiversity not
have 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 not
nonhuman 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 for
A nature-friendly matrix
ARTICLE TOOLS http://science.sciencemag.org/content/362/6412/eaau6020
MATERIALS
SUPPLEMENTARY http://science.sciencemag.org/content/suppl/2018/10/17/362.6412.eaau6020.DC1
CONTENT
RELATED http://science.sciencemag.org/content/sci/362/6412/287.full
REFERENCES http://science.sciencemag.org/content/362/6412/eaau6020#BIBL
This article cites 142 articles, 29 of which you can access for free
PERMISSIONS http://www.sciencemag.org/help/reprints-and-permissions
Terms of ServiceUse of this article is subject to the
is a registered trademark of AAAS.Science
licensee American Association for the Advancement of Science. No claim to original U.S. Government Works. The title
Science, 1200 New York Avenue NW, Washington, DC 20005. 2017 © 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 http://science.sciencemag.org/Downloaded from
www.sciencemag.org/content/362/6412/eaau6020/suppl/DC1
Supplementary Materials for
Landscapes that work for biodiversity and people
C. Kremen* and A. M. Merenlender
*Corresponding author: ckremen@berkeley.edu
Published 19 October 2018, Science 362, eaau6020 (2018)
DOI: 10.1126/science.aau6020
This PDF file includes:
Tables S1 and S2
References
1
Table S1.
Conservation instruments that have potential to promote working lands conservation along with representative examples and
challenges. Voluntary Incentives (red), Market Instruments (blue), Environmental Regulations and Compliance through Offsets
(orange), and Governance (purple).
Conservation instruments
Example applied to working landscapes
Challenges
Conservation easement: A voluntary
legal agreement between a landowner and
a private or public land trust that
permanently limits designated land uses in
order to protect conservation values in
exchange for payment or tax benefits.
The Nature Conservancy is the largest
easement holder in the US.
Approximately 46% of its easement
holdings are on working
landscapes with ranching, forestry, or
farming more likely to be designated as
buffers to enhance biodiversity in the
surrounding area than easements without
these uses (107).
1) Have been amended in the past meaning
perpetuity is uncertain.
2) Hard to monitor compliance and prescriptive
management activities in particular.
3) Private land trusts may prove not to be durable
institutions over time.
4) Can impact public tax revenue.
5) Public funds are often expended by private
institutions without public oversight for the
initial investment choices and oversight (108)
6) May not lead to landscape-level objectives if
not tied to a larger-scale acquisition plan.
Incentive programs for landowners:
public or private funds for land
management activities that can help with
the cost of restoring or maintaining
working landscapes; sometimes matched
by monetary or in-kind contributions from
the landowner.
US Farm Bill Conservation Programs
such as Conservation Reserve Program
and Environmental Quality Incentives
Program fund best management practices
on farmlands, including set-asides,
restoration, and sustainable agriculture
practices.
1) Transaction costs could be high for
government.
2) Adoption rates may be low if application
process is cumbersome or incentives
insufficient.
3) Reversible over time – once contracts are
finished, landowners may fail to renew or
support may no longer be available, which can
lead to fairly extensive land cover change, such
as the loss of grasslands and wetlands in
response to increased corn prices in the US
2
Conservation instruments
Example applied to working landscapes
Challenges
Midwest, causing farmers to let Conservation
Reserve Program contracts lapse(109)
4) Not likely to lead to landscape-level objectives,
unless connected to a landscape-level initiative.
Transfer development rights:
Development is permitted at higher than
allowable densities in exchange for
paying for the lost opportunity costs
associated with reducing development
elsewhere, creating higher density
development and reducing urban sprawl
that can present a threat to working
landscapes; thereby, protecting open
space that can be managed as working
lands.
A) 20,974 ha of agricultural land was
conserved in the densely-developed
Baltimore–Washington, USA, region,
through transferring development rights,
resulting in lower density zoning and
hence reducing the threat from future
development to productive farmland
(110).
B) Land rights trading has achieved
farmland preservation in China leading
the way for future application in
developing countries (111).
1) Loss of open space in higher density urban
areas where access of residents to open space is
already limited.
2) May inflate prices in low density areas thereby
excluding low income residents.
3) Transaction costs mostly borne by private
beneficiaries creating a barrier if the incentive
to increase development is not sufficient to
warrant a trade for protections elsewhere (112)
4) Can be viewed by government as complicated
to implement (113)
Certification and labeling schemes:
Sustainability standards are adopted
voluntarily by landowners and paired with
compliance verification, traceability and
labels, potentially leading to improved
farm, forest or rangeland management
compatible with working lands
conservation
Smithsonian’s “Bird-Friendly” coffee
agroforests; Rainforest Alliance green
frog label (114); organic cacao
agroforestry in Bolivia (115).
1) Limit on environmental standards that can be
met due to costs of implementation and
monitoring beyond what can be passed on to
consumer.
2) Mismatch between spatial targeting of
certification for conservation versus market
goals.
3) Economic benefits are sometimes realized by
end-seller and not by farmer, who may bear
costs (114)
4) Not likely to lead to landscape-level objectives,
unless connected to a landscape-level initiative
Sustainable supply chains: Companies
make voluntary commitments, sometimes
A) The Round-table on Responsible Soy
is a consortium-led group with 3
rd
party
1) This approach may have a large impact due to
the economic power and influence of corporate
3
Conservation instruments
Example applied to working landscapes
Challenges
as part of product certification, to improve
sustainability across their supply chain
that may involve better land/water
management practices and thus may
improve conservation outcomes on
working lands.
verification whose standards include no
natural habitat conversion, protection of
high value conservation areas, and best
management practices regarding soils,
water and agrochemical use (61).
B) Some modest reductions in
deforestation are documented due to
adoption of certification standards in
Indonesia (116)
actors engaging in voluntary actions to improve
sustainability (61), but there is a risk of
“greenwashing”. (114)
2) Opportunities to benefit from certification
without actual conservation outcomes; for
example, the certification of sustainable oil
palm production in Indonesia reduced
deforestation across a limited area, while most
of the certified production comes from areas
that had little remaining forest to start with
(116). Overall the supply chain approach has
not yet arrested deforestation due to
commodity cropping (62).
3) Little data exists to evaluate effectiveness of
standards for achieving specific environmental
outcomes. (61)
4) Considerations noted for certification at the
land-owner level also apply, although when
corporations work with many land-owners in a
region, there is the potential for true landscape-
level approaches.
Payments for ecosystem services: Public
or private compensation to individuals or
communities for activities to promote
specific ecosystem services, often from
cultivated, range or forest lands, which
can include payments for biodiversity
conservation, to reduce the growth in
global greenhouse-gas emissions, and
conserve trees for carbon sequestration
(REDD+).
A) The Regional Integrated Silvopastoral
Ecosystem Management Project in
Nicaragua pays private landowners to
adopt land management practices that
favor biodiversity conservation and
carbon sequestration. Payments are
proportional to the level of management
changes and services provided (117).
B) In Wolong Reserve, China, payments
to local residents and forestry enterprises
to cease fuelwood and timber harvest and
1) Narrow focus on one goal such as C-
sequestration can lead to unintended negative
consequences for other goals; for example
when native forests are replaced by fast
growing plantations to sequester carbon more
rapidly.
2) Monetary rewards can undermine intrinsic
stewardship values or cause perceptions of
entitlement to reward for not performing
environmentally-damaging action
4
Conservation instruments
Example applied to working landscapes
Challenges
prevent forest encroachment through
regular monitoring improved forest cover
and habitat suitability for Giant Panda
(Ailuropoda melanoleuca) (118)
3) Inequitable distribution of benefits within and
among communities.
4) Tradeoffs between enrolling large landowners
versus many landowners.
5) Monitoring for accountability is complex and
costly.
6) Often top-down “one size fits all” programs
that don’t allow landowner knowledge and
creativity to thrive (119)
7) Not likely to lead to landscape-level objectives,
unless connected to a landscape-level initiative
Environmental regulations: Laws
designed to protect the environment by
restricting habitat or species loss,
pollution, or environmental degradation,
thereby protecting natural resources on
working lands.
The Brazilian Forest Code requires
private land-owners to set aside a certain
portion of natural habitat as well as
restore riparian areas and maintain hilltop
forests (120) (see also landscape-level
mitigation, below).
1) Inequitable application of regulations across
physical and social contexts – for example, the
Environmental Protection Agency noted fewer
emission standards violations in Hispanic
communities across the USA than warranted,
demonstrating disparities in the enforcement of
environmental laws (121)
2) Economic impacts of regulations are often
perceived as damaging, creating opposition,
when in fact the actual costs are usually small
and short lived (122).
3) Political will to implement regulations can be
limited when constituents perceive an
economic threat.
Mitigation/habitat and carbon banking:
A habitat conservation area, which can
include working landscapes, where offsets
to compensate for impacts to global
warming, species or ecosystems can be
met for a cost.
A) More than 100 conservation banks had
been approved by the United States Fish
and Wildlife Service in 11 states, covering
some 60 threatened and endangered
species on 790,000 acres (123). Some of
these land banks are working cattle
ranches where grazing is compatible with
1) Restoration can often only partially reach the
desired reference condition and generally takes
many years, meaning the impacts of habitat
loss are incurred for some time before they are
fully mitigated (125, 126).
2) For offsets that are far from the impact sites,
may not be ecologically equivalent (127).
5
Conservation instruments
Example applied to working landscapes
Challenges
maintaining endangered species habitat
(e.g. vernal pool species).
B) 86 out of 1838 projects registered with
the main certifier, Voluntary Carbon
Standard, relate to reforestation efforts in
agriculture and forest landscapes (124).
3) When a single reserve or the same habitat acres
are used to offset impacts for multiple
endangered species, there is no real
additionality for conservation beyond the first
species protected by the bank or site.
4) Mitigation banks are often impacted by
surrounding land use development that will
impact resource conservation within the bank
lands in the future (128).
5) Carbon banks require a stable reservoir of
carbon with consistent year-to-year harvest or
no harvest areas, which is much harder for
small landowners to achieve because their
harvest varies widely over time (129).
6) Contracting offsets requires good governance,
but it is often lacking and therefore limits the
number and quality of projects which explains
why so few forestry projects have been
certified under Kyoto's Clean Development
Mechanism (124).
7) Transactions costs can be high (e.g. measuring,
monitoring, verifying, & enforcing mitigation
results) (124).
Landscape-level mitigation: Identify
mitigation opportunities which support
regional conservation priorities such as
enhancing habitat connectivity across
working lands sometimes include habitat
restoration. This type of off-site
mitigation can be implemented through
in-lieu fees, or prior to the impacts
A) A recent amendment to the Brazilian
Forest Code permits landowners that
develop all of their land to purchase
mitigation credits elsewhere within the
same biome, in order to promote
aggregation of restored or conserved
natural habitat. The application of
landscape-level mitigation in Brazil was
found to reduce total business costs by
1) No net loss of habitat will not be met if
preservation of alternative areas is the basis for
mitigation rather than habitat restoration.
2) Mitigation lands may not be ecologically
equivalent to the impacted areas and
equivalency can be difficult to estimate across
space and time (127).
3) Requires strategic conservation planning to
identify priority investment areas.
6
Conservation instruments
Example applied to working landscapes
Challenges
occurring as is the case for advanced
mitigation.
$19 million per 6-year sugarcane growing
cycle while often supporting more species
and storing more carbon (56).
B) The California Department of
Transportation allocated 7.5 million for
conservation easements on farmland,
equivalent to the land that will be
impacted elsewhere by a light rail project,
thereby advancing large landscape
conservation goals.
Community-based natural resource
management: Management authority is
devolved from government or NGO to
local communities to manage or co-
manage natural resources (often common
pool resources) sustainably and for their
own well-being; including lands inhabited
and managed by indigenous peoples.
A) In Mongolia, numerous instances of
CBNRM have been instigated by
government and NGO agencies to replace
the former traditional communal
management of grazing lands, that was
disrupted in the 1920’s by Mongolia’s
revolution. (130).
B) Indigenous lands make up 20% of the
Brazilian Amazon and serve to protect
these regions from out-
right deforestation.
Indigenous lands occupy a far larger
extent than protected parks in the Amazon
and are often located near the most
vulnerable agricultural frontier areas
(131).
1) Competing interests among different
stakeholders within local communities often
exist.
2) Community governance mechanisms can be
insufficient to implement the management
goals (132).
3) Local expertise and available workforce may
be limited and interventions such as river
restoration treatments can benefit from both
indigenous and more recent scientific
knowledge (133).
4) Without full resolution of land rights and
representation by local people at the highest
levels there can be resistance to full adoption or
engagement of land management efforts (132).
Government lands managed for
production. If policies are put in place
on national forests or rangelands to
implement sustainable management and
protect environmental quality these can
contribute to working lands.
These include IUCN V protected areas --
sustainable management systems aimed at
conserving traditional management
systems and the biodiversity that
sometimes depends on them or on
developing sustainable management
1) Legal and management restrictions on use and
development may not be sufficient or may not
be enforced.
2) Working lands management can be severely
impacted if permissible economic activities on
government lands also include highly
7
Conservation instruments
Example applied to working landscapes
Challenges
systems alongside of biodiversity
conservation (IUCN VI). Lonjsko Polje
Nature Park (IUCN V) in Croatia protects
a traditional pasturing system adapted to a
floodplain ecosystem and maintains an
important reservoir of plant biodiversity,
along with globally endangered species
that depends on grazing (134).
damaging actions such as shale gas
development and mining (135).
3) Difficult to reconcile trade-offs between
improving livelihoods in the near term and
environmental protection and conservation
(42).
4) Restoration and management of these
government lands requires ongoing
intervention particularly in arid landscapes
(136) and because government commitment
and investment can wane.
8
Table S2.
Representative examples showing benefits of diversified farming, ranching and forestry systems
for biodiversity and livelihood outcomes
5
Outcomes
Representative Examples
Biodiversity and connectivity
promoted
1) Organic agriculture enhances biodiversity of many
taxonomic groups relative to conventional agriculture (137–
139).
2) Adding hedgerows comprised of diverse native shrubs and
trees enhances native bee and bird biodiversity in intensively-
farmed landscapes of California relative to unmanaged field
edges (140, 141).
3) Agroforestry provides habitat for tropical forest species like
the mantled howling monkey (Allouatta palliada) and
enhances connectivity between forest reserves (48, 142).
Ecosystem services increased
1) A meta-analysis showed that use of cover crops in place of
winter fallows stored 0.32+ 0.08 Mg C/ha/year, which globally
could compensate for 8% of the annual emissions from
agriculture (143).
2) In tropical coffee systems, the abundance of specific
insectivorous birds doubles pest control of the coffee berry
borer, and is enhanced by increased forest cover both within
and surrounding the farm (avoided damages estimated at
US$75–US$310/ ha/year, 50)
3) Retaining narrow wooded corridors through pasture boosted
pollination success in habitat fragments by 14.3 times,
providing ecosystem function benefits (144).
4) Revegetation with native grass seeds and soil manipulations,
show improved forage across the Great Basin rangelands,
USA, because native perennial grasses provide forage over a
longer season, and increase infiltration of limited rainfall keeps
forage palatable for longer (145).
Negative environmental
externalities minimized
1)By integrating strips of restored prairie onto 10% of the area
of corn and soy bean monocultures in Iowa, phosphorus and
soils entering waterways were reduced by 4.3 and 20-fold,
respectively (86).
2) By planting nectar-rich floral strips along rice fields in Asia
to support natural enemies of crop pests, farmers reduced
insecticide applications by 70% while enhancing yields by 5%
(77).
3) Through complex crop rotations, herbicide use was reduced
by 88% leading to 200-fold reduction in freshwater toxicity,
9
Outcomes
Representative Examples
while increasing yields and maintaining profits (despite higher
labor costs, 73).
Yields increased
1)Planting companion plants to create a push-pull system
controlled two devastating pests of maize (corn-borers and
Striga weeds), tripling yields while also producing fodder for
cattle and improving soil fertility (146).
2) In Bolivia, while cacao yields were lower in experimental
shade agroforestry plots compared to full sun cacao, total yield
of food plants more than doubled in agroforesty plots (147).
3) In a meta-analysis of annual inter-crops, 81% of responses
showed greater yields for the intercrop compared to the sole
crops based on land equivalent ratios (LER); LERs increased
significantly with greater temporal niche differentiation of the
two crops (148).
Profits enhanced/ markets
accessed
1) By reducing input use through diversification practices,
farmers cut costs and enhanced profits (77)
2) Farmers can increase profits by targeting less productive
lands for replacement, either with habitat for beneficial insects
(leading to increased yields on the remaining 92 – 97% of
cropped area, 78), or by planting perennials, which reduced
variable costs (leading to reduced variable costs, 87).
3) Using certification systems for best practices can bring
higher premiums and profits to farmers (e.g., organic, 149),
create market access and green supply chains (e.g. eco-labelled
coffee or cacao, 114).
Food security and livelihoods
enhanced
1) By providing improved fodder for cattle for companion x
plants in push pull maize/sorghum agriculture in Africa, cattle
provide more milk which provides better nutrition and an
income source for women and children (146).
2) Richness and evenness of plants and animals in home
gardens that were managed for consumption were strong
positive predictors of food security during a drought for
subsistence farmers in the Yucatan (150)
3) Farm production diversity led to greater dietary diversity
and improved child health among subsistence farmers in
Malawi (151, 152), although elsewhere, other factors such as
market access affected dietary diversity in important ways
(153).
4) Muyuy’s small farmers near Iquitos, Peru plant more than
260 varieties of plants in agroforestry plots resembling natural
forest, resulting in high agricultural output and immense
diversity of species (154).
Climate adaptation and
mitigation enhanced
1)Cacao farmers perceived that increasing the number and
types of trees in agroforestry plots was a method for adapting
to increased incidence of flood, drought and heat in Bolivia, as
10
Outcomes
Representative Examples
well as disease (115). These agroforestry systems also stored
on average 50 Mg/ha more C than full sun monocultures,
contributing to C-mitigation (155).
2) Corn and soy grown in more complex rotations exhibited
greater yields and more stability during hot and dry periods in
the USA (32), and water infiltration that reduced drought
effects was markedly improved in complex organic rotations
compared to conventional monocultures (156).
4) Large-scale riparian vegetation restoration coupled with
changes in stream flow management could enhance carbon
sequestration benefits (measured as high as 20,915 kg/ha of
biomass and carbon storage near the river compared to close to
zero 2000 m from the river), in Northwest China (157).
Significant contributions
made to global food supply
1) Resource-conserving (agroecological) systems tend to
boost production relative to unimproved (subsistence)
farming methods, on average by 79% (76)
2) Smallholder farms less than 2 ha produce 30 – 34% of the
world’s food by kcal on just 24% of the agricultural land
area (70).
3) About 50% of the world’s smallholder farmers practice
resource-conserving agriculture (158).
4) While difficult to estimate precisely, it follows from the
previous 3 points that agroecological methods currently
contribute a substantial amount to world food production,
and that there is much room for improvement with broader
adoption of these methods.
11
References and Notes
1. J. L. Payne, A. M. Bush, N. A. Heim, M. L. Knope, D. J. McCauley, Ecological selectivity of
the emerging mass extinction in the oceans. Science 353, 1284–1286 (2016).
doi:10.1126/science.aaf2416 Medline
2. G. Ceballos, P. R. Ehrlich, R. Dirzo, Biological annihilation via the ongoing sixth mass
extinction signaled by vertebrate population losses and declines. Proc. Natl. Acad. Sci.
U.S.A. 114, E6089–E6096 (2017). Medline
3. G. Ceballos, P. R. Ehrlich, A. D. Barnosky, A. García, R. M. Pringle, T. M. Palmer,
Accelerated modern human-induced species losses: Entering the sixth mass extinction.
Sci. Adv. 1, e1400253 (2015). doi:10.1126/sciadv.1400253 Medline
4. C. A. Hallmann, M. Sorg, E. Jongejans, H. Siepel, N. Hofland, H. Schwan, W. Stenmans, A.
Müller, H. Sumser, T. Hörren, D. Goulson, H. de Kroon, More than 75 percent decline
over 27 years in total flying insect biomass in protected areas. PLOS ONE 12, e0185809
(2017). doi:10.1371/journal.pone.0185809 Medline
5. T. Newbold, L. N. Hudson, S. L. L. Hill, S. Contu, I. Lysenko, R. A. Senior, L. Börger, D. J.
Bennett, A. Choimes, B. Collen, J. Day, A. De Palma, S. Díaz, S. Echeverria-Londoño,
M. J. Edgar, A. Feldman, M. Garon, M. L. K. Harrison, T. Alhusseini, D. J. Ingram, Y.
Itescu, J. Kattge, V. Kemp, L. Kirkpatrick, M. Kleyer, D. L. P. Correia, C. D. Martin, S.
Meiri, M. Novosolov, Y. Pan, H. R. P. Phillips, D. W. Purves, A. Robinson, J. Simpson,
S. L. Tuck, E. Weiher, H. J. White, R. M. Ewers, G. M. Mace, J. P. W. Scharlemann, A.
Purvis, Global effects of land use on local terrestrial biodiversity. Nature 520, 45–50
(2015). doi:10.1038/nature14324 Medline
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 mountain ranges. Proc. Natl. Acad. Sci. U.S.A. 115, 6004–6009 (2018).
doi:10.1073/pnas.1720141115 Medline
8. R. M. Pringle, Upgrading protected areas to conserve wild biodiversity. Nature 546, 91–99
(2017). doi:10.1038/nature22902 Medline
9. J. M. Halley, N. Monokrousos, A. D. Mazaris, W. D. Newmark, D. Vokou, Dynamics of
extinction debt across five taxonomic groups. Nat. Commun. 7, 12283 (2016).
doi:10.1038/ncomms12283 Medline
10. W. F. Laurance, D. C. Useche, J. Rendeiro, M. Kalka, C. J. A. Bradshaw, S. P. Sloan, S. G.
Laurance, M. Campbell, K. Abernethy, P. Alvarez, V. Arroyo-Rodriguez, P. Ashton, J.
Benítez-Malvido, A. Blom, K. S. Bobo, C. H. Cannon, M. Cao, R. Carroll, C. Chapman,
R. Coates, M. Cords, F. Danielsen, B. De Dijn, E. Dinerstein, M. A. Donnelly, D.
Edwards, F. Edwards, N. Farwig, P. Fashing, P.-M. Forget, M. Foster, G. Gale, D. Harris,
R. Harrison, J. Hart, S. Karpanty, W. J. Kress, J. Krishnaswamy, W. Logsdon, J. Lovett,
W. Magnusson, F. Maisels, A. R. Marshall, D. McClearn, D. Mudappa, M. R. Nielsen, R.
Pearson, N. Pitman, J. van der Ploeg, A. Plumptre, J. Poulsen, M. Quesada, H. Rainey, D.
Robinson, C. Roetgers, F. Rovero, F. Scatena, C. Schulze, D. Sheil, T. Struhsaker, J.
Terborgh, D. Thomas, R. Timm, J. N. Urbina-Cardona, K. Vasudevan, S. J. Wright, J. C.
Arias-G, L. Arroyo, M. Ashton, P. Auzel, D. Babaasa, F. Babweteera, P. Baker, O.
12
Banki, M. Bass, I. Bila-Isia, S. Blake, W. Brockelman, N. Brokaw, C. A. Brühl, S.
Bunyavejchewin, J.-T. Chao, J. Chave, R. Chellam, C. J. Clark, J. Clavijo, R. Congdon,
R. Corlett, H. S. Dattaraja, C. Dave, G. Davies, Bde. M. Beisiegel, Rde. N. da Silva, A.
Di Fiore, A. Diesmos, R. Dirzo, D. Doran-Sheehy, M. Eaton, L. Emmons, A. Estrada, C.
Ewango, L. Fedigan, F. Feer, B. Fruth, J. G. Willis, U. Goodale, S. Goodman, J. C. Guix,
P. Guthiga, W. Haber, K. Hamer, I. Herbinger, J. Hill, Z. Huang, I. F. Sun, K. Ickes, A.
Itoh, N. Ivanauskas, B. Jackes, J. Janovec, D. Janzen, M. Jiangming, C. Jin, T. Jones, H.
Justiniano, E. Kalko, A. Kasangaki, T. Killeen, H. B. King, E. Klop, C. Knott, I. Koné, E.
Kudavidanage, J. L. Ribeiro, J. Lattke, R. Laval, R. Lawton, M. Leal, M. Leighton, M.
Lentino, C. Leonel, J. Lindsell, L. Ling-Ling, K. E. Linsenmair, E. Losos, A. Lugo, J.
Lwanga, A. L. Mack, M. Martins, W. S. McGraw, R. McNab, L. Montag, J. M.
Thompson, J. Nabe-Nielsen, M. Nakagawa, S. Nepal, M. Norconk, V. Novotny, S.
O’Donnell, M. Opiang, P. Ouboter, K. Parker, N. Parthasarathy, K. Pisciotta, D.
Prawiradilaga, C. Pringle, S. Rajathurai, U. Reichard, G. Reinartz, K. Renton, G.
Reynolds, V. Reynolds, E. Riley, M.-O. Rödel, J. Rothman, P. Round, S. Sakai, T.
Sanaiotti, T. Savini, G. Schaab, J. Seidensticker, A. Siaka, M. R. Silman, T. B. Smith, S.
S. de Almeida, N. Sodhi, C. Stanford, K. Stewart, E. Stokes, K. E. Stoner, R. Sukumar,
M. Surbeck, M. Tobler, T. Tscharntke, A. Turkalo, G. Umapathy, M. van Weerd, J. V.
Rivera, M. Venkataraman, L. Venn, C. Verea, C. V. de Castilho, M. Waltert, B. Wang,
D. Watts, W. Weber, P. West, D. Whitacre, K. Whitney, D. Wilkie, S. Williams, D. D.
Wright, P. Wright, L. Xiankai, P. Yonzon, F. Zamzani, Averting biodiversity collapse in
tropical forest protected areas. Nature 489, 290–294 (2012). doi:10.1038/nature11318
Medline
11. M. C. Urban, Climate change. Accelerating extinction risk from climate change. Science 348,
571–573 (2015). doi:10.1126/science.aaa4984 Medline
12. S. R. Loarie, P. B. Duffy, H. Hamilton, G. P. Asner, C. B. Field, D. D. Ackerly, The velocity
of climate change. Nature 462, 1052–1055 (2009). doi:10.1038/nature08649 Medline
13. L. O. Frishkoff, D. S. Karp, L. K. M’Gonigle, C. D. Mendenhall, J. Zook, C. Kremen, E. A.
Hadly, G. C. Daily, Loss of avian phylogenetic diversity in neotropical agricultural
systems. Science 345, 1343–1346 (2014). doi:10.1126/science.1254610 Medline
14. G. Chapron, P. Kaczensky, J. D. C. Linnell, M. von Arx, D. Huber, H. Andrén, J. V. López-
Bao, M. Adamec, F. Álvares, O. Anders, L. Balčiauskas, V. Balys, P. Bedő, F. Bego, J.
C. Blanco, U. Breitenmoser, H. Brøseth, L. Bufka, R. Bunikyte, P. Ciucci, A. Dutsov, T.
Engleder, C. Fuxjäger, C. Groff, K. Holmala, B. Hoxha, Y. Iliopoulos, O. Ionescu, J.
Jeremić, K. Jerina, G. Kluth, F. Knauer, I. Kojola, I. Kos, M. Krofel, J. Kubala, S.
Kunovac, J. Kusak, M. Kutal, O. Liberg, A. Majić, P. Männil, R. Manz, E. Marboutin, F.
Marucco, D. Melovski, K. Mersini, Y. Mertzanis, R. W. Mysłajek, S. Nowak, J. Odden,
J. Ozolins, G. Palomero, M. Paunović, J. Persson, H. Potočnik, P.-Y. Quenette, G. Rauer,
I. Reinhardt, R. Rigg, A. Ryser, V. Salvatori, T. Skrbinšek, A. Stojanov, J. E. Swenson,
L. Szemethy, A. Trajçe, E. Tsingarska-Sedefcheva, M. Váňa, R. Veeroja, P. Wabakken,
M. Wölfl, S. Wölfl, F. Zimmermann, D. Zlatanova, L. Boitani, Recovery of large
carnivores in Europe’s modern human-dominated landscapes. Science 346, 1517–1519
(2014). doi:10.1126/science.1257553 Medline
13
15. M. D. Reynolds, B. L. Sullivan, E. Hallstein, S. Matsumoto, S. Kelling, M. Merrifield, D.
Fink, A. Johnston, W. M. Hochachka, N. E. Bruns, M. E. Reiter, S. Veloz, C. Hickey, N.
Elliott, L. Martin, J. W. Fitzpatrick, P. Spraycar, G. H. Golet, C. McColl, C. Low, S. A.
Morrison, Dynamic conservation for migratory species. Sci. Adv. 3, e1700707 (2017).
doi:10.1126/sciadv.1700707 Medline
16. C. Kremen, Reframing the land-sparing/land-sharing debate for biodiversity conservation.
Ann. N.Y. Acad. Sci. 1355, 52–76 (2015). doi:10.1111/nyas.12845 Medline
17. C. D. Mendenhall, A. Shields-Estrada, A. J. Krishnaswami, G. C. Daily, Quantifying and
sustaining biodiversity in tropical agricultural landscapes. Proc. Natl. Acad. Sci. U.S.A.
113, 14544–14551 (2016). doi:10.1073/pnas.1604981113 Medline
18. C. D. Mendenhall, D. S. Karp, C. F. J. Meyer, E. A. Hadly, G. C. Daily, Predicting
biodiversity change and averting collapse in agricultural landscapes. Nature 509, 213–
217 (2014). doi:10.1038/nature13139 Medline
19. C. M. Kennedy, E. H. C. Grant, M. C. Neel, W. F. Fagan, P. P. Marra, Landscape matrix
mediates occupancy dynamics of Neotropical avian insectivores. Ecol. Appl. 21, 1837–
1850 (2011). doi:10.1890/10-1044.1 Medline
20. J. A. Foley, R. Defries, G. P. Asner, C. Barford, G. Bonan, S. R. Carpenter, F. S. Chapin, M.
T. Coe, G. C. Daily, H. K. Gibbs, J. H. Helkowski, T. Holloway, E. A. Howard, C. J.
Kucharik, C. Monfreda, J. A. Patz, I. C. Prentice, N. Ramankutty, P. K. Snyder, Global
consequences of land use. Science 309, 570–574 (2005). doi:10.1126/science.1111772
Medline
21. N. Ramankutty, Z. Mehrabi, K. Waha, L. Jarvis, C. Kremen, M. Herrero, L. H. Rieseberg,
Trends in global agricultural land use: Implications for environmental health and food
security. Annu. Rev. Plant Biol. 69, 789–815 (2018). doi:10.1146/annurev-arplant-
042817-040256 Medline
22. F. Isbell, A. Gonzalez, M. Loreau, J. Cowles, S. Díaz, A. Hector, G. M. Mace, D. A. Wardle,
M. I. O’Connor, J. E. Duffy, L. A. Turnbull, P. L. Thompson, A. Larigauderie, Linking
the influence and dependence of people on biodiversity across scales. Nature 546, 65–72
(2017). doi:10.1038/nature22899 Medline
23. F. Isbell, P. R. Adler, N. Eisenhauer, D. Fornara, K. Kimmel, C. Kremen, D. K. Letourneau,
M. Liebman, H. W. Polley, S. Quijas, M. Scherer-Lorenzen, Benefits of increasing plant
diversity in sustainable agroecosystems. J. Ecol. 105, 871–879 (2017). doi:10.1111/1365-
2745.12789
24. D. J. Eastburn, A. T. O’Geen, K. W. Tate, L. M. Roche, Multiple ecosystem services in a
working landscape. PLOS ONE 12, e0166595 (2017). doi:10.1371/journal.pone.0166595
Medline
25. D. A. Driscoll, S. C. Banks, P. S. Barton, D. B. Lindenmayer, A. L. Smith, Conceptual
domain of the matrix in fragmented landscapes. Trends Ecol. Evol. 28, 605–613 (2013).
doi:10.1016/j.tree.2013.06.010 Medline
26. W. F. Fagan, E. E. Holmes, Quantifying the extinction vortex. Ecol. Lett. 9, 51–60 (2006).
Medline
14
27. F. S. Chapin 3rd, S. R. Carpenter, G. P. Kofinas, C. Folke, N. Abel, W. C. Clark, P. Olsson,
D. M. S. Smith, B. Walker, O. R. Young, F. Berkes, R. Biggs, J. M. Grove, R. L. Naylor,
E. Pinkerton, W. Steffen, F. J. Swanson, Ecosystem stewardship: Sustainability strategies
for a rapidly changing planet. Trends Ecol. Evol. 25, 241–249 (2010).
doi:10.1016/j.tree.2009.10.008 Medline
28. F. Isbell, V. Calcagno, A. Hector, J. Connolly, W. S. Harpole, P. B. Reich, M. Scherer-
Lorenzen, B. Schmid, D. Tilman, J. van Ruijven, A. Weigelt, B. J. Wilsey, E. S. Zavaleta,
M. Loreau, High plant diversity is needed to maintain ecosystem services. Nature 477,
199–202 (2011). doi:10.1038/nature10282 Medline
29. M. G. E. Mitchell, E. M. Bennett, A. Gonzalez, Linking landscape connectivity and
ecosystem service provision: Current knowledge and research gaps. Ecosystems 16, 894–
908 (2013). doi:10.1007/s10021-013-9647-2
30. M. Loreau, N. Mouquet, A. Gonzalez, Biodiversity as spatial insurance in heterogeneous
landscapes. Proc. Natl. Acad. Sci. U.S.A. 100, 12765–12770 (2003).
doi:10.1073/pnas.2235465100 Medline
31. C. Kremen, A. Miles, Ecosystem services in biologically diversified versus conventional
farming systems: Benefits, externalities, and trade-offs. Ecol. Soc. 17, 40 (2012).
doi:10.5751/ES-05035-170440
32. A. C. M. Gaudin, T. N. Tolhurst, A. P. Ker, K. Janovicek, C. Tortora, R. C. Martin, W. Deen,
Increasing crop diversity mitigates weather variations and improves yield stability. PLOS
ONE 10, e0113261 (2015). doi:10.1371/journal.pone.0113261 Medline
33. S. M. Philpott, B. B. Lin, S. Jha, S. J. Brines, A multi-scale assessment of hurricane impacts
on agricultural landscapes based on land use and topographic features. Agric. Ecosyst.
Environ. 128, 12–20 (2008). doi:10.1016/j.agee.2008.04.016
34. L. C. Ponisio, L. K. M’Gonigle, K. C. Mace, J. Palomino, P. de Valpine, C. Kremen,
Diversification practices reduce organic to conventional yield gap. Proc. R. Soc. London
Ser. B 282, 20141396 (2015). doi:10.1098/rspb.2014.1396 Medline
35. R. J. Hobbs, E. Higgs, C. M. Hall, P. Bridgewater, F. S. Chapin III, E. C. Ellis, J. J. Ewel, L.
M. Hallett, J. Harris, K. B. Hulvey, S. T. Jackson, P. L. Kennedy, C. Kueffer, L. Lach, T.
C. Lantz, A. E. Lugo, J. Mascaro, S. D. Murphy, C. R. Nelson, M. P. Perring, D. M.
Richardson, T. R. Seastedt, R. J. Standish, B. M. Starzomski, K. N. Suding, P. M.
Tognetti, L. Yakob, L. Yung, Managing the whole landscape: Historical, hybrid, and
novel ecosystems. Front. Ecol. Environ. 12, 557–564 (2014). doi:10.1890/130300
36. N. Estrada-Carmona, A. K. Hart, F. A. J. DeClerck, C. A. Harvey, J. C. Milder, Integrated
landscape management for agriculture, rural livelihoods, and ecosystem conservation: An
assessment of experience from Latin America and the Caribbean. Landsc. Urban Plan.
129, 1–11 (2014). doi:10.1016/j.landurbplan.2014.05.001
37. H. Gu, S. M. Subramanian, Drivers of change in socio-ecological production landscapes:
Implications for better management. Ecol. Soc. 19, 41 (2014). doi:10.5751/ES-06283-
190141
38. M. K. Anderson, Tending the Wild: Native American Knowledge and the Management of
California’s Natural Resources (Univ. of California Press, 2005).
15
39. E. M. Bignal, D. I. McCracken, The nature conservation value of European traditional
farming systems. Environ. Rev. 8, 149–171 (2000). doi:10.1139/a00-009
40. S. K. Lowder, J. Skoet, T. Raney, The number, size, and distribution of farms, smallholder
farms, and family farms worldwide. World Dev. 87, 16–29 (2016).
doi:10.1016/j.worlddev.2015.10.041
41. C. L. Gray, S. L. L. Hill, T. Newbold, L. N. Hudson, L. Börger, S. Contu, A. J. Hoskins, S.
Ferrier, A. Purvis, J. P. W. Scharlemann, Local biodiversity is higher inside than outside
terrestrial protected areas worldwide. Nat. Commun. 7, 12306 (2016).
doi:10.1038/ncomms12306 Medline
42. C. L. Shafer, Cautionary thoughts on IUCN protected area management categories V-VI.
Global Ecol. Conserv. 3, 331–348 (2015). doi:10.1016/j.gecco.2014.12.007
43. F. A. J. DeClerck, R. Chazdon, K. D. Holl, J. C. Milder, B. Finegan, A. Martinez-Salinas, P.
Imbach, L. Canet, Z. Ramos, Biodiversity conservation in human-modified landscapes of
Mesoamerica: Past, present and future. Biol. Conserv. 143, 2301–2313 (2010).
doi:10.1016/j.biocon.2010.03.026
44. C. A. Harvey, A. Medina, D. M. Sánchez, S. Vílchez, B. Hernández, J. C. Saenz, J. M. Maes,
F. Casanoves, F. L. Sinclair, Patterns of animal diversity in different forms of tree cover
in agricultural landscapes. Ecol. Appl. 16, 1986–1999 (2006). doi:10.1890/1051-
0761(2006)016[1986:POADID]2.0.CO;2 Medline
45. Ç. H. Şekercioğlu, S. R. Loarie, F. Oviedo-Brenes, C. D. Mendenhall, G. C. Daily, P. R.
Ehrlich, Tropical countryside riparian corridors provide critical habitat and connectivity
for seed-dispersing forest birds in a fragmented landscape. J. Ornithol. 156 (Suppl. 1),
343–353 (2015). doi:10.1007/s10336-015-1299-x
46. C. A. Harvey, C. Villanueva, J. Villacís, M. Chacón, D. Muñoz, M. López, M. Ibrahim, R.
Gómez, R. Taylor, J. Martinez, A. Navas, J. Saenz, D. Sánchez, A. Medina, S. Vilchez,
B. Hernández, A. Perez, F. Ruiz, F. López, I. Lang, F. L. Sinclair, Contribution of live
fences to the ecological integrity of agricultural landscapes. Agric. Ecosyst. Environ. 111,
200–230 (2005). doi:10.1016/j.agee.2005.06.011
47. A. Medina, C. A. Harvey, D. S. Merlo, S. Vílchez, B. Hernández, Bat diversity and
movement in an agricultural landscape in Matiguás, Nicaragua. Biotropica 39, 120–128
(2007). doi:10.1111/j.1744-7429.2006.00240.x
48. K. Williams-Guillen, C. McCann, J. C. Martinez Sanchez, F. Koontz, Resource availability
and habitat use by mantled howling monkeys in a Nicaraguan coffee plantation: Can
agroforests serve as core habitat for a forest mammal? Anim. Conserv. 9, 331–338
(2006). doi:10.1111/j.1469-1795.2006.00042.x
49. J. Avelino, A. Romero-Gurdián, H. F. Cruz-Cuellar, F. A. J. Declerck, Landscape context
and scale differentially impact coffee leaf rust, coffee berry borer, and coffee root-knot
nematodes. Ecol. Appl. 22, 584–596 (2012). doi:10.1890/11-0869.1 Medline
50. D. S. Karp, C. D. Mendenhall, R. F. Sandí, N. Chaumont, P. R. Ehrlich, E. A. Hadly, G. C.
Daily, Forest bolsters bird abundance, pest control and coffee yield. Ecol. Lett. 16, 1339–
1347 (2013). doi:10.1111/ele.12173 Medline
16
51. C. A. Harvey, M. Chacón, C. I. Donatti, E. Garen, L. Hannah, A. Andrade, L. Bede, D.
Brown, A. Calle, J. Chará, C. Clement, E. Gray, M. H. Hoang, P. Minang, A. M.
Rodríguez, C. Seeberg-Elverfeldt, B. Semroc, S. Shames, S. Smukler, E. Somarriba, E.
Torquebiau, J. van Etten, E. Wollenberg, Climate-smart landscapes: Opportunities and
challenges for integrating adaptation and mitigation in tropical agriculture. Conserv. Lett.
7, 77–90 (2014). doi:10.1111/conl.12066
52. A. T. H. Keeley, G. Basson, D. R. Cameron, N. E. Heller, P. R. Huber, C. A. Schloss, J. H.
Thorne, A. M. Merenlender, Making habitat connectivity a reality. Conserv. Biol. (2018).
doi:10.1111/cobi.13158 Medline
53. W. C. Morse, J. L. Schedlbauer, S. E. Sesnie, B. Finegan, C. A. Harvey, S. J. Hollenhorst, K.
L. Kavanagh, D. Stoian, J. D. Wulfhorst, Consequences of environmental service
payments for forest retention and recruitment in a Costa Rican biological corridor. Ecol.
Soc. 14, 23 (2009). doi:10.5751/ES-02688-140123
54. J. Owley, D. Takacs, “Flexible Conservation in Uncertain Times,” in Contemporary Issues in
Climate Change Law and Policy: Essays Inspired by the IPCC (UC Hastings Research
Paper 180, Univ. at Buffalo School of Law, 2016), pp. 65–102;
https://papers.ssrn.com/sol3/papers.cfm?abstract_id=2700658.
55. J. Alvez, A. L. Schmitt Filho, J. Farley, G. Alarcon, A. C. Fantini, The potential for
agroecosystems to restore ecological corridors and sustain farmer livelihoods: Evidence
from Brazil. Ecol. Restor. 30, 288–290 (2012). doi:10.3368/er.30.4.288
56. C. M. Kennedy, D. A. Miteva, L. Baumgarten, P. L. Hawthorne, K. Sochi, S. Polasky, J. R.
Oakleaf, E. M. Uhlhorn, J. Kiesecker, Bigger is better: Improved nature conservation and
economic returns from landscape-level mitigation. Sci. Adv. 2, e1501021 (2016).
doi:10.1126/sciadv.1501021 Medline
57. I. Scoones, P. Newell, M. Leach, in The Politics of Green Transformations, I. Scoones, M.
Leach, P. Newell, Eds. (Earthscan from Routledge, 2015), pp. 1–24.
58. M. B. Mascia, M. Mills, When conservation goes viral: The diffusion of innovative
biodiversity conservation policies and practices. Conserv. Lett. 11, e12442 (2018).
doi:10.1111/conl.12442
59. D. F. Doak, V. J. Bakker, B. E. Goldstein, B. Hale, What is the future of conservation?
Trends Ecol. Evol. 29, 77–81 (2014). doi:10.1016/j.tree.2013.10.013 Medline
60. K. Hamrick, “State of private investment in conservation 2016: A landscape assessment of an
emerging market” (Forest Trends, 2016).
61. R. Chaplin-Kramer, M. Jonell, A. Guerry, E. F. Lambin, A. J. Morgan, D. Pennington, N.
Smith, J. A. Franch, S. Polasky, Ecosystem service information to benefit sustainability
standards for commodity supply chains. Ann. N.Y. Acad. Sci. 1355, 77–97 (2015).
doi:10.1111/nyas.12961 Medline
62. P. G. Curtis, C. M. Slay, N. L. Harris, A. Tyukavina, M. C. Hansen, Classifying drivers of
global forest loss. Science 361, 1108–1111 (2018). doi:10.1126/science.aau3445 Medline
17
63. D. Pannell, G. R. Marshall, N. Barr, A. Curtis, F. Vanclay, R. Wilkinson, Understanding and
promoting adoption of conservation practices by rural landholders. Aust. J. Exp. Agric.
46, 1407–1424 (2006). doi:10.1071/EA05037
64. M. Leach, I. Scoones, in The Politics of Green Transformations, I. Scoones, P. Newell, M.
Leach, Eds. (Earthscan from Routledge, 2015), pp. 119–133.
65. D. R. Cameron, D. C. Marvin, J. M. Remucal, M. C. Passero, Ecosystem management and
land conservation can substantially contribute to California’s climate mitigation goals.
Proc. Natl. Acad. Sci. U.S.A. 114, 12833–12838 (2017). doi:10.1073/pnas.1707811114
Medline
66. N. Ramankutty, A. T. Evan, C. Monfreda, J. A. Foley, Farming the planet: 1. Geographic
distribution of global agricultural lands in the year 2000. Global Biogeochem. Cycles 22,
GB1003 (2008). doi:10.1029/2007GB002952
67. M. A. Altieri, The ecological role of biodiversity in agroecosystems. Agric. Ecosyst. Environ.
74, 19–31 (1999). doi:10.1016/S0167-8809(99)00028-6
68. International Panel of Experts on Sustainable Food Systems (IPES-Food), “From uniformity
to diversity: A paradigm shift from industrial agriculture to diversified agroecological
systems” (IPES-Food, 2016); http://www.ipes-
food.org/images/Reports/UniformityToDiversity_FullReport.pdf.
69. C. Kremen, A. Iles, C. M. Bacon, Diversified farming systems: An agro-ecological, systems-
based alternative to modern industrial agriculture. Ecol. Soc. 17, 44 (2012).
doi:10.5751/ES-05103-170444
70. V. Ricciardi, N. Ramankutty, Z. Mehrabi, L. Jarvis, B. Chookolingo, How much of the
world’s food do smallholders produce? Global Food Sec. 17, 64–72 (2018).
doi:10.1016/j.gfs.2018.05.002
71. M. Montenegro, Banking on wild relatives to feed the world. Gastron. J. Crit. Food Stud. 16,
1–8 (2016). doi:10.1525/gfc.2016.16.1.1
72. A. Iles, R. Marsh, Nurturing diversified farming systems in industrialized countries: How
public policy can contribute. Ecol. Soc. 17, 42 (2012). doi:10.5751/ES-05041-170442
73. A. S. Davis, J. D. Hill, C. A. Chase, A. M. Johanns, M. Liebman, Increasing cropping system
diversity balances productivity, profitability and environmental health. PLOS ONE 7,
e47149 (2012). doi:10.1371/journal.pone.0047149 Medline
74. B. Phalan, M. Onial, A. Balmford, R. E. Green, Reconciling food production and biodiversity
conservation: Land sharing and land sparing compared. Science 333, 1289–1291 (2011).
doi:10.1126/science.1208742 Medline
75. K. Garbach, J. C. Milder, F. A. J. DeClerck, M. Montenegro de Wit, L. Driscoll, B. Gemmill-
Herren, Examining multi-functionality for crop yield and ecosystem services in five
systems of agroecological intensification. Int. J. Agric. Sustain. 15, 11–28 (2017).
doi:10.1080/14735903.2016.1174810
76. J. Pretty, Z. P. Bharucha, Sustainable intensification in agricultural systems. Ann. Bot. 114,
1571–1596 (2014). doi:10.1093/aob/mcu205 Medline
18
77. G. M. Gurr, Z. Lu, X. Zheng, H. Xu, P. Zhu, G. Chen, X. Yao, J. Cheng, Z. Zhu, J. L.
Catindig, S. Villareal, H. Van Chien, Q. Cuong, C. Channoo, N. Chengwattana, L. P.
Lan, H. Hai, J. Chaiwong, H. I. Nicol, D. J. Perovic, S. D. Wratten, K. L. Heong, Multi-
country evidence that crop diversification promotes ecological intensification of
agriculture. Nat. Plants 2, 16014 (2016). doi:10.1038/nplants.2016.14 Medline
78. R. F. Pywell, M. S. Heard, B. A. Woodcock, S. Hinsley, L. Ridding, M. Nowakowski, J. M.
Bullock, Wildlife-friendly farming increases crop yield: Evidence for ecological
intensification. Proc. R. Soc. London Ser. B 282, 20151740 (2015).
doi:10.1098/rspb.2015.1740 Medline
79. L. Li, S.-M. Li, J.-H. Sun, L.-L. Zhou, X.-G. Bao, H.-G. Zhang, F.-S. Zhang, Diversity
enhances agricultural productivity via rhizosphere phosphorus facilitation on phosphorus-
deficient soils. Proc. Natl. Acad. Sci. U.S.A. 104, 11192–11196 (2007).
doi:10.1073/pnas.0704591104 Medline
80. D. K. Letourneau, I. Armbrecht, B. S. Rivera, J. M. Lerma, E. J. Carmona, M. C. Daza, S.
Escobar, V. Galindo, C. Gutiérrez, S. D. López, J. L. Mejía, A. M. A. Rangel, J. H.
Rangel, L. Rivera, C. A. Saavedra, A. M. Torres, A. R. Trujillo, Does plant diversity
benefit agroecosystems? A synthetic review. Ecol. Appl. 21, 9–21 (2011).
doi:10.1890/09-2026.1 Medline
81. Y. Zhu, H. Chen, J. Fan, Y. Wang, Y. Li, J. Chen, J. Fan, S. Yang, L. Hu, H. Leung, T. W.
Mew, P. S. Teng, Z. Wang, C. C. Mundt, Genetic diversity and disease control in rice.
Nature 406, 718–722 (2000). doi:10.1038/35021046 Medline
82. S. F. Bender, C. Wagg, M. G. A. van der Heijden, An underground revolution: Biodiversity
and soil ecological engineering for agricultural sustainability. Trends Ecol. Evol. 31,
440–452 (2016). doi:10.1016/j.tree.2016.02.016 Medline
83. I. Perfecto, J. Vandermeer, A. Wright, Nature’s Matrix: Linking Agriculture, Conservation
and Food Sovereignty (Earthscan, 2009).
84. J. A. Hilty, A. M. Merenlender, Use of riparian corridors and vineyards by mammalian
predators in northern California. Conserv. Biol. 18, 126–135 (2004). doi:10.1111/j.1523-
1739.2004.00225.x
85. W. E. Thogmartin, L. López-Hoffman, J. Rohweder, J. Diffendorfer, R. Drum, D. Semmens,
S. Black, I. Caldwell, D. Cotter, P. Drobney, L. L. Jackson, M. Gale, D. Helmers, S.
Hilburger, E. Howard, K. Oberhauser, J. Pleasants, B. Semmens, O. Taylor, P. Ward, J.
F. Weltzin, R. Wiederholt, Restoring monarch butterfly habitat in the Midwestern US:
‘All hands on deck.’ Environ. Res. Lett. 12, 074005 (2017). doi:10.1088/1748-
9326/aa7637
86. L. A. Schulte, J. Niemi, M. J. Helmers, M. Liebman, J. G. Arbuckle, D. E. James, R. K.
Kolka, M. E. O’Neal, M. D. Tomer, J. C. Tyndall, H. Asbjornsen, P. Drobney, J. Neal, G.
Van Ryswyk, C. Witte, Prairie strips improve biodiversity and the delivery of multiple
ecosystem services from corn-soybean croplands. Proc. Natl. Acad. Sci. U.S.A. 114,
11247–11252 (2017). doi:10.1073/pnas.1620229114 Medline
87. E. Brandes, G. S. McNunn, L. A. Schulte, I. J. Bonner, D. J. Muth, B. A. Babcock, B.
Sharma, E. A. Heaton, Subfield profitability analysis reveals an economic case for
19
cropland diversification. Environ. Res. Lett. 11, 014009 (2016). doi:10.1088/1748-
9326/11/1/014009
88. O. De Schutter, “Agroecology and the right to food,” report presented at the 16th Session of
the United Nations Human Rights Council [A/HRC/16/49], Geneva, Switzerland, 8
March 2011.
89. K. G. MacDicken, P. Sola, J. E. Hall, C. Sabogal, M. Tadoum, C. de Wasseige, Global
progress toward sustainable forest management. For. Ecol. Manage. 352, 47–56 (2015).
doi:10.1016/j.foreco.2015.02.005
90. H. G. Lund, Accounting for the world’s rangelands. Soc. Range Manage. 29, 3–10 (2007).
91. S. Gauthier, M.-A. Vaillancourt, D. Kneeshaw, P. Drapeau, L. De Grandpré, Y. Claveau, D.
Paré, in Ecosystem Management in the Boreal Forest, S. Gauthier, M.-A. Vaillancourt,
A. Leduc, L. De Grandpré, D. Kneeshaw, H. Morin, P. Drapeau, Y. Bergeron, Eds.
(Univ. du Québec, 2009), pp. 13–38.
92. M. Mangel, L. M. Talbot, G. K. Meffe, M. T. Agardy, D. L. Alverson, J. Barlow, D. B.
Botkin, G. Budowski, T. Clark, J. Cooke, R. H. Crozier, P. K. Dayton, D. L. Elder, C. W.
Fowler, S. Funtowicz, J. Giske, R. J. Hofman, S. J. Holt, S. R. Kellert, L. A. Kimball, D.
Ludgwig, K. Magnusson, B. S. Malayang III, C. Mann, E. A. Norse, S. P. Northridge, W.
F. Perrin, C. Perrings, E. A. Norse, S. P. Northridge, W. F. Perrin, C. Perrings, R. M.
Peterman, G. B. Rabb, H. A. Regier, J. E. Reynolds III, K. Sherman, M. P. Sissenwine, T.
D. Smith, A. Starfield, R. J. Taylor, M. F. Tillman, C. Toft, J. R. Twiss Jr., J. Wilen, T. P.
Young, Principles for the conservation of wild living resources. Ecol. Appl. 6, 338–362
(1996). doi:10.2307/2269369
93. J. A. Layzer, Natural Experiments: Ecosystem-Based Management and the Environment
(MIT, 2008).
94. P. Campos, L. Huntsinger, J. L. Oviedo, P. F. Starrs, M. Diaz, R. B. Standiford, G. Montero,
Eds., Mediterranean Oak Woodland Working Landscapes: Dehesas of Spain and
Ranchlands of California (Springer Science+Business Media, 2013).
95. J. A. Ramírez, M. Díaz, The role of temporal shrub encroachment for the maintenance of
Spanish holm oak Quercus ilex dehesas. For. Ecol. Manage. 255, 1976–1983 (2008).
doi:10.1016/j.foreco.2007.12.019
96. M. A. Xenopoulos, D. M. Lodge, J. Alcamo, M. Marker, K. Schulze, D. P. Van Vuuren,
Scenarios of freshwater fish extinctions from climate change and water withdrawal.
Global Change Biol. 11, 1557–1564 (2005). doi:10.1111/j.1365-2486.2005.001008.x
97. S. D. Polade, A. Gershunov, D. R. Cayan, M. D. Dettinger, D. W. Pierce, Precipitation in a
warming world: Assessing projected hydro-climate changes in California and other
Mediterranean climate regions. Sci. Rep. 7, 10783 (2017). doi:10.1038/s41598-017-
11285-y Medline
98. F. R. Hauer, H. Locke, V. J. Dreitz, M. Hebblewhite, W. H. Lowe, C. C. Muhlfeld, C. R.
Nelson, M. F. Proctor, S. B. Rood, Gravel-bed river floodplains are the ecological nexus
of glaciated mountain landscapes. Sci. Adv. 2, e1600026 (2016).
doi:10.1126/sciadv.1600026 Medline
20
99. A. T. Keeley, D. D. Ackerly, D. R. Cameron, N. E. Heller, P. R. Huber, C. A. Schloss, J. H.
Thorne, A. M. Merenlender, New concepts, models, and assessments of climate-wise
connectivity. Environ. Res. Lett. 13, 073002 (2018). doi:10.1088/1748-9326/aacb85
100. W. Sutherland, L. V. Dicks, N. Ockendon, R. Smith, Eds., What Works in Conservation
(Open Book, ed. 2, 2017), vol. 2.
101. J. Bongaarts, S. W. Sinding, A response to critics of family planning programs. Int.
Perspect. Sex. Reprod. Health 35, 39–44 (2009). doi:10.1363/3503909 Medline
102. E. W. Sanderson, J. Walston, J. G. Robinson, From bottleneck to breakthrough:
Urbanization and the future of biodiversity conservation. Bioscience 68, 412–426 (2018).
doi:10.1093/biosci/biy039 Medline
103. C. J. Bradshaw, B. W. Brook, Human population reduction is not a quick fix for
environmental problems. Proc. Natl. Acad. Sci. U.S.A. 111, 16610–16615 (2014).
doi:10.1073/pnas.1410465111 Medline
104. J. Poore, T. Nemecek, Reducing food’s environmental impacts through producers and
consumers. Science 360, 987–992 (2018). doi:10.1126/science.aaq0216 Medline
105. Great Eastern Ranges, www.ger.org.au.
106. V. Athreya, M. Odden, J. D. C. Linnell, J. Krishnaswamy, U. Karanth, Big cats in our
backyards: Persistence of large carnivores in a human dominated landscape in India.
PLOS ONE 8, e57872 (2013). doi:10.1371/journal.pone.0057872 Medline
107. A. R. Rissman, L. Lozier, T. Comendant, P. Kareiva, J. M. Kiesecker, M. R. Shaw, A. M.
Merenlender, Conservation easements: Biodiversity protection and private use. Conserv.
Biol. 21, 709–718 (2007). doi:10.1111/j.1523-1739.2007.00660.x Medline
108. A. A. M. Merenlender, L. Huntsinger, G. Guthey, S. K. Fairfax, Land trusts and
conservation easements : Who is conserving what for whom? Conserv. Biol. 18, 65–76
(2004). doi:10.1111/j.1523-1739.2004.00401.x
109. C. K. Wright, M. C. Wimberly, Recent land use change in the Western Corn Belt threatens
grasslands and wetlands. Proc. Natl. Acad. Sci. U.S.A. 110, 4134–4139 (2013).
doi:10.1073/pnas.1215404110 Medline
110. R. Pruetz, N. Standridge, What makes transfer of development rights work?: Success factors
from research and practice. J. Am. Plann. Assoc. 75, 78–87 (2008).
doi:10.1080/01944360802565627
111. H. Wang, R. Tao, L. Wang, F. Su, Farmland preservation and land development rights
trading in Zhejiang, China. Habitat Int. 34, 454–463 (2010).
doi:10.1016/j.habitatint.2009.12.004
112. S. Shahab, J. P. Clinch, E. O’Neill, Estimates of transaction costs in transfer of development
rights programs. J. Am. Plann. Assoc. 84, 61–75 (2018).
doi:10.1080/01944363.2017.1406816
113. A. J. Shriar, A. Akins, Transfer of development rights, growth management, and landscape
conservation in Virginia. Local Environ. 23, 1–19 (2018).
doi:10.1080/13549839.2017.1382458
21
114. T. Tscharntke, J. C. Milder, G. Schroth, Y. Clough, F. DeClerck, A. Waldron, R. Rice, J.
Ghazoul, Conserving biodiversity through certification of tropical agroforestry crops at
local and landscape scales. Conserv. Lett. 8, 14–23 (2015). doi:10.1111/conl.12110
115. J. Jacobi, M. Schneider, P. Bottazzi, M. Pillco, P. Calizaya, S. Rist, Agroecosystem
resilience and farmers’ perceptions of climate change impacts on cocoa farms in Alto
Beni, Bolivia. Renew. Agric. Food Syst. 30, 170–183 (2015).
doi:10.1017/S174217051300029X
116. K. M. Carlson, R. Heilmayr, H. K. Gibbs, P. Noojipady, D. N. Burns, D. C. Morton, N. F.
Walker, G. D. Paoli, C. Kremen, Effect of oil palm sustainability certification on
deforestation and fire in Indonesia. Proc. Natl. Acad. Sci. U.S.A. 115, 121–126 (2018).
doi:10.1073/pnas.1704728114 Medline
117. S. Pagiola, A. R. Rios, A. Arcenas, Can the poor participate in payments for environmental
services? Lessons from the Silvopastoral Project in Nicaragua. Environ. Dev. Econ. 13,
299–325 (2008). doi:10.1017/S1355770X08004270
118. M.-N. Tuanmu, A. Viña, W. Yang, X. Chen, A. M. Shortridge, J. Liu, Effects of payments
for ecosystem services on wildlife habitat recovery. Conserv. Biol. 30, 827–835 (2016).
doi:10.1111/cobi.12669 Medline
119. K. M. A. Chan, E. Anderson, M. Chapman, K. Jespersen, P. Olmsted, Payments for
ecosystem services: Rife with problems and potential—for transformation towards
sustainability. Ecol. Econ. 140, 110–122 (2017). doi:10.1016/j.ecolecon.2017.04.029
120. B. Soares-Filho, R. Rajão, M. Macedo, A. Carneiro, W. Costa, M. Coe, H. Rodrigues, A.
Alencar, Land use. Cracking Brazil’s forest code. Science 344, 363–364 (2014).
doi:10.1126/science.1246663 Medline
121. D. M. Konisky, C. Reenock, Regulatory enforcement, riskscapes, and environmental
justice. Policy Stud. J. 46, 7–36 (2018). doi:10.1111/psj.12203
122. A. Dechezleprêtre, M. Sato, The impacts of environmental regulations on competitiveness.
Rev. Environ. Econ. Policy 11, 183–206 (2017). doi:10.1093/reep/rex013
123. L. Scarlett, R. Espanchin-Niell, M. McKinney, The Endangered Species Act at 40: New
tools for conservation. Resources 184, 20–27 (2013).
124. G. C. van Kooten, Forest carbon offsets and carbon emissions trading: Problems of
contracting. For. Policy Econ. 75, 83–88 (2017). doi:10.1016/j.forpol.2016.12.006
125. J. M. Bullock, J. Aronson, A. C. Newton, R. F. Pywell, J. M. Rey-Benayas, Restoration of
ecosystem services and biodiversity: Conflicts and opportunities. Trends Ecol. Evol. 26,
541–549 (2011). doi:10.1016/j.tree.2011.06.011 Medline
126. N. Goldberg, K. C. Reiss, Accounting for wetland loss: Wetland mitigation trends in
northeast Florida 2006–2013. Wetlands 36, 373–384 (2016). doi:10.1007/s13157-016-
0749-4
127. F. Quétier, S. Lavorel, Assessing ecological equivalence in biodiversity offset schemes:
Key issues and solutions. Biol. Conserv. 144, 2991–2999 (2011).
doi:10.1016/j.biocon.2011.09.002
22
128. H. Levrel, P. Scemama, A.-C. Vaissiere, Should we be wary of mitigation banking?
Evidence regarding the risks associated with this wetland offset arrangement in Florida.
Ecol. Econ. 135, 136–149 (2017). doi:10.1016/j.ecolecon.2016.12.025
129. H. Bigsby, Carbon banking: Creating flexibility for forest owners. For. Ecol. Manage. 257,
378–383 (2009). doi:10.1016/j.foreco.2008.09.018
130. M. E. Fernández-Giménez, B. Batkhishig, B. Batbuyan, T. Ulambayar, Lessons from the
dzud: Community-based rangeland management increases the adaptive capacity of
Mongolian herders to winter disasters. World Dev. 68, 48–65 (2015).
doi:10.1016/j.worlddev.2014.11.015
131. D. Nepstad, S. Schwartzman, B. Bamberger, M. Santilli, D. Ray, P. Schlesinger, P.
Lefebvre, A. Alencar, E. Prinz, G. Fiske, A. Rolla, Inhibition of Amazon deforestation
and fire by parks and indigenous lands. Conserv. Biol. 20, 65–73 (2006).
doi:10.1111/j.1523-1739.2006.00351.x Medline
132. R. S. Wilson, T. Pearce, K. Jones, S. Fleischfresser, B. Davis, G. Jones, S. Lieske,
Indigenous land management in peri-urban landscapes: An Australian example. Soc. Nat.
Resour. 31, 335–350 (2018). doi:10.1080/08941920.2017.1383543
133. B. D. Hartman, D. A. Cleveland, O. A. Chadwick, Linking changes in knowledge and
attitudes with successful land restoration in indigenous communities. Restor. Ecol. 24,
749–760 (2016). doi:10.1111/rec.12347
134. G. Gugić, D. Župan, I. Zupan, in Protected Landscapes and Wild Biodiversity, N. Dudley,
S. Stolton, Eds. (Values of Protected Landscapes and Seascapes 3, International Union
for Conservation of Nature and Natural Resources, 2012), pp. 19–26.
135. D. Costa, J. Jesus, D. Branco, A. Danko, A. Fiúza, Extensive review of shale gas
environmental impacts from scientific literature (2010–2015). Environ. Sci. Pollut. Res.
Int. 24, 14579–14594 (2017). doi:10.1007/s11356-017-8970-0 Medline
136. R. J. Hobbs, Where to from here? Challenges for restoration and revegetation in a fast-
changing world. Rangeland J. 39, 563–566 (2017). doi:10.1071/RJ17053
137. C. Winqvist, J. Bengtsson, T. Aavik, F. Berendse, L. W. Clement, S. Eggers, C. Fischer, A.
Flohre, F. Geiger, J. Liira, T. Pärt, C. Thies, T. Tscharntke, W. W. Weisser, R.
Bommarco, Mixed effects of organic farming and landscape complexity on farmland
biodiversity and biological control potential across Europe. J. Appl. Ecol. 48, 570–579
(2011). doi:10.1111/j.1365-2664.2010.01950.x
138. D. Gabriel, S. M. Sait, J. A. Hodgson, U. Schmutz, W. E. Kunin, T. G. Benton, Scale
matters: The impact of organic farming on biodiversity at different spatial scales. Ecol.
Lett. 13, 858–869 (2010). doi:10.1111/j.1461-0248.2010.01481.x Medline
139. J. Bengtsson, J. Ahnström, A. C. Weibull, The effects of organic agriculture on biodiversity
and abundance: A meta-analysis. J. Appl. Ecol. 42, 261–269 (2005). doi:10.1111/j.1365-
2664.2005.01005.x
140. L. C. Ponisio, L. K. M’Gonigle, C. Kremen, On-farm habitat restoration counters biotic
homogenization in intensively managed agriculture. Global Change Biol. 22, 704–715
(2016). doi:10.1111/gcb.13117 Medline
23
141. S. K. Heath, C. U. Soykan, K. L. Velas, R. Kelsey, S. M. Kross, A bustle in the hedgerow:
Woody field margins boost on farm avian diversity and abundance in an intensive
agricultural landscape. Biol. Conserv. 212, 153–161 (2017).
doi:10.1016/j.biocon.2017.05.031
142. S. A. Bhagwat, K. J. Willis, H. J. B. Birks, R. J. Whittaker, Agroforestry: A refuge for
tropical biodiversity? Trends Ecol. Evol. 23, 261–267 (2008).
doi:10.1016/j.tree.2008.01.005 Medline
143. C. Poeplau, A. Don, Carbon sequestration in agricultural soils via cultivation of cover crops
– A meta-analysis. Agric. Ecosyst. Environ. 200, 33–41 (2015).
doi:10.1016/j.agee.2014.10.024
144. U. Kormann, C. Scherber, T. Tscharntke, N. Klein, M. Larbig, J. J. Valente, A. S. Hadley,
M. G. Betts, Corridors restore animal-mediated pollination in fragmented tropical forest
landscapes. Proc. R. Soc. London Ser. B 283, 20152347 (2016).
doi:10.1098/rspb.2015.2347
145. D. S. Pilliod, J. L. Welty, G. R. Toevs, Seventy-five years of vegetation treatments on
public rangelands in the Great Basin of North America. Rangelands 39, 1–9 (2017).
doi:10.1016/j.rala.2016.12.001
146. Z. Khan, C. Midega, J. Pittchar, J. Pickett, T. Bruce, Push–pull technology: A conservation
agriculture approach for integrated management of insect pests, weeds and soil health in
Africa. Int. J. Agric. Sustain. 9, 162–170 (2011). doi:10.3763/ijas.2010.0558
147. M. Schneider, C. Andres, G. Trujillo, F. Alcon, P. Amurrio, E. Perez, F. Weibel, J. Milz,
Cocoa and total system yields of organic and conventional agroforestry vs. monoculture
systems in a long-term field trial in Bolivia. Exp. Agric. 53, 351–374 (2017).
doi:10.1017/S0014479716000417
148. Y. Yu, T. J. Stomph, D. Makowski, W. van der Werf, Temporal niche differentiation
increases the land equivalent ratio of annual intercrops: A meta-analysis. Field Crops
Res. 184, 133–144 (2015). doi:10.1016/j.fcr.2015.09.010
149. D. W. Crowder, J. P. Reganold, Financial competitiveness of organic agriculture on a global
scale. Proc. Natl. Acad. Sci. U.S.A. 112, 7611–7616 (2015).
doi:10.1073/pnas.1423674112 Medline
150. D. Sampson, thesis, University of California, Santa Cruz (2015).
151. S. L. Dwivedi, E. T. Lammerts van Bueren, S. Ceccarelli, S. Grando, H. D. Upadhyaya, R.
Ortiz, Diversifying food systems in the pursuit of sustainable food production and healthy
diets. Trends Plant Sci. 22, 842–856 (2017). doi:10.1016/j.tplants.2017.06.011 Medline
152. A. D. Jones, On-farm crop species richness is associated with household diet diversity and
quality in subsistence- and market-oriented farming households in Malawi. J. Nutr. 147,
86–96 (2017). doi:10.3945/jn.116.235879 Medline
153. K. T. Sibhatu, V. V. Krishna, M. Qaim, Production diversity and dietary diversity in
smallholder farm households. Proc. Natl. Acad. Sci. U.S.A. 112, 10657–10662 (2015).
doi:10.1073/pnas.1510982112 Medline
24
154. M. Pinedo-Vasquez, J. Barletti Pasqualle, D. Del Castillo Torres, K. Coffey, A tradition of
change: The dynamic relationship between biodiversity and society in sector Muyuy,
Peru. Environ. Sci. Policy 5, 43–53 (2002). doi:10.1016/S1462-9011(02)00023-0
155. J. Jacobi, C. Andres, M. Schneider, M. Pillco, P. Calizaya, S. Rist, Carbon stocks, tree
diversity, and the role of organic certification in different cocoa production systems in
Alto Beni, Bolivia. Agrofor. Syst. 88, 1117–1132 (2014). doi:10.1007/s10457-013-9643-
8
156. D. W. Lotter, R. Seidel, W. Liebhardt, The performance of organic and conventional
cropping systems in an extreme climate year. Am. J. Altern. Agric. 18, 146–154 (2003).
doi:10.1079/AJAA200345
157. T. Aishan, F. Betz, U. Halik, B. Cyffka, A. Rouzi, Biomass carbon sequestration potential
by riparian forest in the Tarim River watershed, Northwest China: Implication for the
mitigation of climate change impact. Forests 9, 196 (2018). doi:10.3390/f9040196
158. M. A. Altieri, V. M. Toledo, The agroecological revolution in Latin America: Rescuing
nature, ensuring food sovereignty and empowering peasants. J. Peasant Stud. 38, 587–
612 (2011). doi:10.1080/03066150.2011.582947