Article
Securing Nature’s Contrib
utions to People requires
at least 20%–25% (semi-)natural habitat in human-
modified landscapes
Graphical abstract
Highlights
d We assess habitat quantity, quality, and spatial configuration

in human-modified landscapes

d At least 20%–25% habitat per km2 is needed to sustain

nature’s contributions to people

d Only one-third of global human-modified lands meet this

minimum level

d Local actions should be adopted based on community needs,

knowledge, and capacities
Mohamed et al., 2024, One Earth 7, 59–71
January 19, 2024 ª 2023 The Authors. Published by Elsevier Inc
https://doi.org/10.1016/j.oneear.2023.12.008
Authors

Awaz Mohamed, Fabrice DeClerck,

Peter H. Verburg, ..., Sarah K. Jones,

Ina C. Meier, Ben Stewart-Koster

Correspondence
awaz.mohamed2@gmail.com

In brief

Biodiversity loss threatens crucial human

well-being aspects, including food

production, water quality, climate

regulation, and recreation. We assess the

minimum level of (semi-)natural habitat in

agricultural and urban areas to sustain

these benefits. We find that below 20%–

25% (semi-)natural habitat per km2, the

supply of these benefits significantly

declines. Alarmingly, two-thirds of global

urban and agricultural lands fall below

this level. Our study offers a broad target

for conservation efforts beyond natural

areas to enhance human well-being.
.
ll

mailto:awaz.mohamed2@gmail.�com
https://doi.org/10.1016/j.oneear.2023.12.008
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OPEN ACCESS

ll
Article

Securing Nature’s Contributions to People requires
at least 20%–25% (semi-)natural habitat
in human-modified landscapes
Awaz Mohamed,1,12,13,* Fabrice DeClerck,2 Peter H. Verburg,3,4 David Obura,5 Jesse F. Abrams,6 Noelia Zafra-Calvo,7

JuanRocha,8,9 Natalia Estrada-Carmona,2 Alexander Fremier,10 Sarah K. Jones,2 Ina C.Meier,1 andBen Stewart-Koster11
1Functional Forest Ecology, Universit€at Hamburg, 22885 Barsb€uttel, Germany
2Alliance of Biodiversity International and CIAT, CGIAR, Montpellier, France
3Institute for Environmental Studies, Vrije Universiteit Amsterdam, Amsterdam, the Netherlands
4Swiss Federal Institute for Forest, Snow and Landscape Research (WSL), Birmensdorf, Switzerland
5CORDIO East Africa, Mombasa, Kenya
6Global Systems Institute, University of Exeter, Exeter EX4 4QE, UK
7Basque Centre for Climate Change bc3, Scientific Campus of the University of the Basque Country, Biscay, Spain
8Future Earth Secretariat, Stockholm, Sweden
9Stockholm Resilience Centre, Stockholm University, Stockholm, Sweden
10School of the Environment, Washington State University, Pullman, WA 99164, USA
11Australian Rivers Institute, Griffith University, 170 Kessels Road, Nathan, QLD 4111, Australia
12X (formerly Twitter): @awazmsm
13Lead contact

*Correspondence: awaz.mohamed2@gmail.com
https://doi.org/10.1016/j.oneear.2023.12.008
SCIENCE FOR SOCIETY Biodiversity is rapidly declining, affecting benefits critical to human well-being,
including food production, water quality, climate regulation, and recreation spaces. This decline is partic-
ularly challenging in urban and agricultural areas significantly modified by humans but where reliance on
biodiversity’s benefits is nevertheless high. We assess the minimum level of (semi-)natural habitat needed
in human-modified landscapes to support the supply of these benefits. We find that biodiversity’s capacity
to pollinate crops, regulate pests and diseases, maintain clear water, and limit soil erosion significantly de-
clines when habitat area falls below 20%–25% per km2. This same limit applies in urban areas to maintain
recreation spaces for people. We find that approximately two-thirds of agricultural and urban areas globally
fall below this level. This broad target can be used in urban and agricultural areas tomanage and regenerate
ecosystem functions to enhance human well-being.
SUMMARY
The cascading effects of biodiversity decline on human well-being present a pressing challenge for sustain-
able development. Conservation efforts often prioritize safeguarding specific species, habitats, or intact eco-
systems but overlook biodiversity’s fundamental role in providing Nature’s Contributions to People (NCP) in
human-modified landscapes. Here, we systematically review 154 peer-reviewed studies to estimate the min-
imum levels of (semi-)natural habitat quantity, quality, and spatial configuration needed in human-modified
landscapes to secure functional integrity essential for sustaining NCP provision. We find that the provision
of multiple NCP is threatened when (semi-)natural habitat in the landscape falls below an area of 20%–
25% for each km2. Five NCP almost completely disappear below a level of 10% habitat. The exact quantity,
quality, and spatial configuration of habitat required depends on local context and specific NCP. Today,
about two-thirds of human-modified lands have insufficient (semi-)natural habitat, requiring action for NCP
regeneration. Our findings serve as a generic guideline to target conservation actions outside natural areas.
INTRODUCTION
 human well-being.1 Such contributions range from climate, wa-
Recent global assessments demonstrate a clear decline in living

nature and its contributions to people with cascading effects on
One Earth 7, 59–71, J
This is an open access article und
ter, and nutrient cycle regulation at global and regional scales

to pollination, pest control, and physical and psychological ex-

periences at local scales. These benefits are generally referred
anuary 19, 2024 ª 2023 The Authors. Published by Elsevier Inc. 59
er the CC BY license (http://creativecommons.org/licenses/by/4.0/).

https://twitter.com/awazmsm
mailto:awaz.mohamed2@gmail.com
https://doi.org/10.1016/j.oneear.2023.12.008
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ll
OPEN ACCESS Article
to as ecosystem services or Nature’s Contributions to People

(NCP) and comprise the ecosystem functions that directly or

indirectly contribute to human well-being and quality of life.2

Local-scale NCP are particularly important in human-modified

landscapes due to the intensive interaction between human pop-

ulations and natural ecosystems, often having a high number of

beneficiaries and a greater potential for the use of NCP. These

areas, however, are often ignored in global-scale studies inform-

ing conservation priorities that tend to focus on intact natural

lands and wilderness areas.3

Biodiversity has multiple facets, including genes, species,

populations, evolutionary history, ecosystem functions, and

contributions to people, as well as a variety of social and cultural

dimensions. Most attention in biodiversity conservation is given

to halting the conversion of remaining intact natural ecosystems,

protecting the unique species they hold,4,5 and the important

contributions they make to Earth system functioning (goal A of

the Kunming-Montreal Global Biodiversity Framework). These

are critically important conservation objectives; however,

(semi-)natural habitats in human-modified lands and waters are

often overlooked in conservation policies and global target

setting, despite the critical roles they play in supporting human

well-being6 as well as in conserving biodiversity.7 Human-modi-

fied lands cover approximately 50% of the ice-free terrestrial

land area and range from urban to agricultural areas.8 The signif-

icant decline of ecosystem functions and contributions to people

in such areas is incompatible with several of the Sustainable

Development Goals and the agreed targets of the Kunming-

Montreal Global Biodiversity Framework, notably Target 10 on

sustainable production.9 However, we currently lack generaliz-

able and operational metrics describing the functions of biodi-

versity embedded within human-modified lands and the mini-

mum level of biodiversity in these landscapes needed to

support human well-being.10,11 Identifying such metrics is chal-

lenged by the highly context-specific conditions under which

biodiversity supports multiple ecosystem processes in human-

modified landscapes, making it also challenging to define syn-

thetic policy objectives.10

A functional integrity metric has been proposed to capture the

multiple dimensions and interactions between species and the

environment in a synthetic measure,12,13 but clear evidence of

the minimum level of functional integrity required remains

missing.14 In this study, we refer to functional integrity as the ca-

pacity of the ecosystem to contribute to biosphere processes

and to sustain multiple NCP provision through the presence of

ecologically functional communities of species. It addresses

both Earth-system-scale biosphere regulation processes and

landscape-scale provisioning of local NCP. Functional integrity

complements biodiversity metrics used in conservation biology

by recognizing the important NCP that can be provided by natu-

ral vegetation but also by altered (non-native or non-intact) vege-

tation in agricultural, urban, and other human-modified areas.

Our reference to a minimum level of functional integrity refers

to a level of ecosystem function below which there is a substan-

tial risk of experiencing a strong decline in NCP provision, jeop-

ardizing the well-being of those dependent on these NCP. This

minimum level should not be interpreted as the amount required

for sufficient NCP provision, nor as an estimate of NCP supply,

as both can strongly vary depending on the local context.
60 One Earth 7, 59–71, January 19, 2024
NCP provision is dependent on the quantity, quality, and

spatial configuration of available (semi-)natural habitat (hereafter

‘‘habitat’’) within the landscape, which can be used as a proxy

measure of functional integrity.15 Habitat quantity refers to the

proportion of (semi-)natural elements present in a landscape.

Habitat quality is a measure of the ability of a habitat to host

and maintain species required for specific ecological functions

and services. The structure and composition of a habitat are

strong determinants of its quality.16 The spatial configuration of

habitat in the landscape influences landscape connectivity and

the distribution of NCP-providing organisms. This includes

both the proximity to habitat and the location of habitat that sup-

ports NCP provision. Adjacency is an important element of NCP

provisioning, notably in managed lands where distances be-

tween habitat (NCP source) and crops or people (NCP benefi-

ciary) can vary. Habitat location within a landscape is also impor-

tant for regulating water quality, particularly in riparian zones,

whereas the distance to source habitat determines access by

mobile NCP-providing organisms. These include pollinators

and pest and disease-controlling organisms where the foraging

range from the home habitat determines themaximum linear dis-

tance for NCP provision. Adjacency also applies to experiences,

where physical access or a reasonable distance from human

residence determines its potential use. The combination of

quantity, quality, and spatial configuration of habitat collectively

underpins functional integrity.

The required habitat quantity, quality, and spatial configura-

tion for NCP provision are strongly context dependent and differ

depending on the NCP, landscape type, and the taxa

involved.17–19 Shorter linear distances from source habitat (a

few hundred meters) and higher connectivity have an important

positive impact on mobile pollinators and pest regulator diver-

sity.18 For NCP provided by sessile or low-mobility functional

groups (e.g., soil erosion control, capture of non-point-source

pollutants from surface and subsurface water, or natural hazards

mitigation), habitat location is extremely important. For example,

sediment and nutrient capture are significantly improved through

vegetation buffers along both sides of waterways, in particular

on stream headwaters.20 Likewise, habitat strategically located

in targeted landscape positions can significantly reduce the fre-

quency, risk, and impact of natural hazards such as shallow

landslides, floods, and soil erosion. Habitat in urban ecosys-

tems, in the form of greenspaces and parks, can provide impor-

tant NCP such as physically and psychologically beneficial expe-

riences that contribute significantly to well-being.21

Numerous ecological studies have studied aspects of the rela-

tionship between habitat quantity, quality, and spatial configura-

tion and the provisioning of NCP. Although these studies confirm

the high context specificity and variability of such relationships,

they consistently indicate that, below certain levels of habitat

quantity, quality, and spatial configuration, NCP provisioning

strongly declines or is even no longer provided.22–26 Studies on

pollination and pest control suggest required levels of 10%–

20% habitat per km2, often based on expert judgment, valid in

a specific land-use or landscape types.15,27,28 To our knowl-

edge, a synthesis of minimum levels for functional integrity

across several NCP and across a wide range of landscapes

has not been conducted to date. Such a synthesis would serve

as a generally applicable guide and provide an overview of the



Figure 1. Minimum quantity of habitat

required for provisioning of each NCP

The lower and upper red lines correspond to the

whiskers (minimum and maximum, respectively),

which indicate the range of the data. The middle red

line represents the median, while the red dots

represent the weighted mean value. The violin

shape indicates kernel density estimation based on

the number of original papers included in the meta-

analyses/reviews reporting the given value. Wider

sections of the violin plot represent a larger number

of papers underlying the given value; the thinner

sections represent a lower body of evidence. All the

values are weighted by the number of papers.

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OPEN ACCESSArticle
order of magnitude of (semi-)natural habitat required in human-

modified landscapes. It also contributes to more effective and

targeted conservation and restoration strategies, promoting sus-

tainable NCP provisioning and safeguarding the well-being of

people reliant on these NCP.

Here, we attempt to determine the minimum quantity, quality,

and spatial configuration needs of (semi-)natural habitat in hu-

man-modified landscapes. These minimum needs indicate the

minimum level of functional integrity essential for maintaining

NCP provision. To achieve this, we conducted a systematic re-

view of the literature, analyzing 74 quantitative peer-reviewed re-

views and 80 narrative studies, comprising in total 4,277 original

studies. We identify the level below which six critical NCP signif-

icantly decline. Our key findings, therefore, delineate the critical

habitat levels necessary for maintaining the following NCP provi-

sion: (1) pollination, (2) pest and disease control, (3) water quality

regulation, (4) soil erosion control, (5) natural hazards mitigation,

and (6) physical and psychological beneficial experiences for

individuals that spend time in natural environments (hereafter

‘‘experiences’’). Our analysis indicates that the capacity of hu-

man-modified landscapes to provide NCP significantly declines

below 20%–25% of habitat per km2, with five NCP provisions

almost completely disappearing below the 10% habitat level.

Currently, about two-thirds of global human-modified land-

scapes lack sufficient area of (semi-)natural habitat within the

landscape to secure this minimum level of NCP provision. Our

proposed levels serve as a general guideline for prioritizing con-

servation initiatives and formulating adaptive, scalable policies

beyond natural areas.

RESULTS

Methods summary
We have selected five regulating and one non-material NCP that

are related to biodiversity and ecosystem functions in different

ways while directly affecting local well-being. These include wa-

ter quality regulation, soil erosion control, crop pollination, pest
control, and benefits for human health.

We use a systematic literature review pro-

tocol of the peer-reviewed literature to

quantitatively synthesize the evidence on

theminimal conditions in terms of quantity,

quality, and spatial configuration of (semi-)

natural habitat necessary for the provision
of the six aforementioned NCP in highly transformed human-

modified landscapes (see ‘‘experimental procedures’’ section

for detailed methods). Minimum values required refer to the level

under which the NCP show a sharp decline or reach very low

values. The median among these observed minimum values is

used to determine the level of (semi-)natural habitat area mini-

mally needed within the landscape. This level is assumed to

secure the multiple ecological functions that underlie the

selected NCP, irrespective of the existence of demand for those

NCP. All values used in establishing the minimum conditions

required are weighted by the number of papers included in

each analyzed review or meta-analysis and represent the me-

dian for each NCP. The values range for individual NCP reflects

variations between studies, whereas the values range across

all NCP represents the variation in median values from the

different NCP.

Habitat quantity
Within the body of literature collected, we coded publications

based on their findings concerning the minimum amount of

(semi-)natural habitat needed within the landscape to secure

six critical local NCP. A total of 94 synthetic and original

studies, encompassing 2,125 original studies, reported relevant

information. Our review concluded that at least 20% habitat is

needed to support pollination and pest and disease control,

with a range of 10%–50% for pollination, and 10%–38% for

pest and disease control, depending on the context. For expe-

riences provided by green spaces, at least 25% habitat is

required, ranging between 19% and 30% depending on the

context (Figure 1; Table 1). Given the dominance of urban

studies for this NCP, our minimum quantity value might be un-

representative of non-urban areas due to variations in transpor-

tation options and alternatives to urban green spaces such as

surrounding croplands.

To protect soil from water-based erosion, at least 50% habitat

at the landscape level is required, with a range of 30%–63% for

specific contexts, depending on slope angle, rainfall intensity,
One Earth 7, 59–71, January 19, 2024 61



Table 1. Estimates of habitat levels for NCP provision

NCP

Taxonomic

groups cited

Minimum habitat

quantity (% km�2)

[range]

Maximum distance

(m)/or position [range] Landscape elements needed

Pollination insects 20% (mean: 21% ± 1%)

[10%–50%]

(total: 172 studies)

<500 m (mean: 989 ± 43 m)

[15–2,000 m]

(total: 288 studies)

rich, diverse habitat with native and

non-native species (floral strips,

floral field margins, floral understory cover;

grassy and woody margins of fields,

hedgerows, woody or silvo-arable corridors

between fields; forest edges and patches

surrounding grassland and shrublands

patches)

Pest and

disease

control

insects, birds,

arachnids

20% (mean: 19% ± 0.2%)

[10%–38%]

(total: 260 studies)

<500 m (mean: 606 ± 23 m)

[10–2,000 m]

(total: 207 studies)

complex habitat with a diverse range of

native species (forest edges and patches;

floral strips, floral field margins, floral

understory cover; grassland, pasture, and

shrubland surrounding patches; grassy and

woody hedgerows and field margins;

woody corridors between fields with

floral understory)

Experiences plants,

birds

25% (mean: 25% ± 0.6%)

[19%–30%]

(total: 26 studies)

<300 m (mean: 311 ± 7 m)

[300–500 m]

(total: 45 studies)

diverse, rich (semi-)natural green spaces

(streets trees canopy cover, public parks,

zoos, gardens, woody and grassy parks,

meadows)

Soil erosion

control

plants 50% (mean: 44% ± 0.6%)

[30%–63%]

(total: 251 studies)

evenly distributed at

the landscape scale

diverse, rich (semi-)natural vegetation cover

(zoned grassy and woody buffers; tree

canopy cover; ground cover with dense

fibrous roots; cover crops such

as grasses and legumes; agroforestry and

woody and grassy hedgerows; mixed

forest, shrublands and grasslands cover;

extensive vegetation management with

inter-row cover or crop cover, no-till

farming, organic farms)

Water quality

regulation

plants 6% (mean: 6% ± 0.1%)

[1.2%–15%]

(total: 1,480 studies)

both sides of streams diverse (semi)-natural vegetative buffers or

strips with diverse range of native species

(three zoned buffers [native forest, shrubs,

and grasses]; forested or mixed forested

and grassy buffers; grassy buffers or

mixed buffers; wetland)

Natural hazards

mitigation

plants 50% (mean: 50.5%)

(total: two studies)

landslides: slope

base or slope bottom

(semi-)natural vegetation cover with diverse

native species (native strong deep-rooted

trees and shrubs with more reinforcing

effect and low surcharge [low height and

low diameter]; spaced young exotic

species [18–20 m] such as popular and

willows; natural young trees;

mixed plantation)

Values constraining the provisioning of the NCP for habitat quantity, quality, and spatial configuration are indicated as levels and represent the median.

All the values are weighted by the number of studies included in the review studies analyzed. The total number of studies refers to the total number of

primary studies considered in articles, reviews, and meta-analyses.

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OPEN ACCESS Article
and landscape type. Regulating stream water quality from

non-point-source pollutants requires a buffer of approximately

28m inwidth on each side of streams. Considering global stream

densities, this minimum buffer width, on average, would corre-

spond to approximately 6% habitat per km2 (Figure 1; Table 1).

The total quantity of habitat needed for specific water quality

functions ranges between 1.2% and 15% depending on the

function in question (nutrient, sediment or pesticide interception

and capture), slope angle, and stream density.
62 One Earth 7, 59–71, January 19, 2024
Identifying the quantity of habitat for reducing landslide risk

(natural hazards mitigation) is more challenging, with environ-

mental variables (geology, slope geometry, soil type, precipita-

tion event frequency, intensity, and duration) often overriding

biological ones (vegetation presence) (Figure 1; Table 1). We

found two studies proposing a quantitative estimate for

regulating landslide risk, advising a minimum of 50% and

60% permanent vegetative cover on steeply sloped lands

(>35�), respectively.29,30



Figure 2. Landscape elements required for provisioning of each NCP

The stacked bar chart showing the proportion of papers recommending specific landscape elements categories that improve habitat quality and support the

provisioning of each NCP. Each bar in the chart represents a whole weighted number of papers analyzed for each NCP (in parentheses), and segments in the bar

with different colors represent different landscape element categories. Natrual habitat: NH; (semi-)natural habitat: SNH.

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OPEN ACCESSArticle
Habitat quality
In our survey, we found 136 synthetic and original papers (en-

compassing 3,755 original studies in total) recommending a

measure of habitat quality in their findings, often expressed as

the range of landscape elements required to support the under-

lying ecological function. We identified six categories of land-

scape elements for enhancing NCP provision: (1) complex

diverse (semi)-natural habitat, (2) complex diverse natural

habitat, (3) diverse floral resources, (4) forest, (5) grassy ele-

ments, and (6) woody elements (Figure 2; Table S1). Natural hab-

itats are areas that have not been significantly modified by hu-

man activities and retain a high level of biodiversity and

ecological integrity. Semi-natural habitats, on the other hand,

are areas that have been modified to some extent by human ac-

tivities but still have many ecological processes intact.1 Diverse

floral resources encompass habitats rich in flowering plants,

while grassy elements are dominated by grasses and herba-

ceous plants. Forests indicate a minimum land area of 0.05–

1.0 ha with tree cover of 10%–30% or more, featuring trees

capable of reaching 2–5 m height at maturity, either in closed

or open forest formations, whereas woody elements comprise

shrubs and trees contributing to landscape diversity and

structural complexity (e.g., shelterbelts, hedgerows, and street

trees).31 The range of landscape elements reported in the re-

viewed literature can take various forms, including strips,

patches, hedgerows, field margins, field borders, ground cover,

canopy cover, and buffers, and, in urban areas, gardens, zoos,

and parks. The quality of habitat required varies depending on

the specific NCP (Figure 2). Nevertheless, Figure 2 illustrates

that 79% of studies we reviewed indicated heterogeneous land-

scapes consisting of complex, diverse (semi-)natural habitat as

themost suitable for supporting multiple NCP provision. Figure 2

also indicates that pollinators demonstrated a notable inclination

toward thriving within rich floristic habitat, particularly those

incorporating wild and native species. In contrast, pest and dis-

ease control organisms tend to be more abundant in complex,

diverse (semi-)natural habitats dominated by diverse woody or

grassy elements rather than being determined by floral resource
availability (Figure 2). For experiences in urban areas, structurally

complex diverse (semi-)natural vegetation including street

trees, public parks, and green spaces are most often habitats

mentioned in the reviewed studies.

The evidence gathered from our study indicates that, to pre-

vent particle detachment driving soil erosion and to intercept de-

tached soil particles transported by water erosion, a structurally

complex, diverse (semi-)natural vegetation cover (encompass-

ing both permanent canopy and ground covers) is required.

This encompasses vegetated buffers, woody and grassy hedge-

rows or agroforests, ground cover or understory vegetation, in-

ter-row vegetated strips, or crop cover with grasses or legumes,

with a dominance of forest and woody elements.

Our reviewed papers indicate that structurally complex and

highly diverse riparian buffers (e.g., zoned buffers consisting of

grassy, shrub, and woody elements), including native species

with diverse root structures, especially when combined with

high stem density, are an important means of slowing excesswa-

ter flows and intercepting detached soil particles (sediment), pes-

ticides, and nutrients from adjacent fields (Figure 2). For steep

slopes, multiple studies emphasize that deep-rooted perennial

cover from diversified fast-growing plantings and understory

vegetation are most effective in reducing landslides.29,32,33

Spatial configuration
We identified 39 synthetic and original papers (representing 415

studies in total) reporting findings on habitat placement within

the landscape and the requiredmaximum linear distance for mo-

bile organisms to access resources from their home habitat. For

pollination and pest and disease control, notably by insects, our

study indicates for both organisms (based on 288 individual

studies for pollination and 207 studies for pest and disease con-

trol) a maximum linear foraging distance of 500 m (ranging be-

tween <0 and 2,000 m for specific taxa within each NCP, under-

lying the variations observed in different studies) from their host

habitat to the target crop field (Figure 3; Table 1). For experiences

obtained in urban ecosystems, most analyzed studies (45

studies in total) indicate 300 m as a maximum reasonable
One Earth 7, 59–71, January 19, 2024 63



Figure 3. Maximum linear distance values be-

tween habitat and beneficiaries (inmeters) for

each NCP

The lower red line and the top red line correspond to

the whiskers (minimum andmaximum, respectively),

which indicate the range of the data. The middle

red line represents the median, while the red dots

represent theweightedmean value. The violin shape

indicates kernel density estimation based on the

number of original papers included in the meta-an-

alyses/reviews reporting the given value. Wider

sections of the violin plot represent a larger number

of papers underlying the given value; the thinner

sections represent a lower body of evidence. All the

values are weighted by the number of papers.

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OPEN ACCESS Article
distance for people to access green spaces based on the iden-

tified positive health impacts of experiencing at least >120min of

nature exposure per week21,34 (Figure 3). We identified from the

collective evidence 300 m (for citizens) to 500 m (for pollination

and pest and disease control, representing the most limiting

median value among these NCP) as the maximum distance

between habitat and target beneficiaries (Figure 3). These dis-

tances represent the minimum conditions required in terms of

spatial configuration; beyond these levels, the NCP provision de-

clines significantly or becomes almost completely absent.

Our review indicates that a complex and diverse vegetation

cover that encompasses at least 50% of the land, with an even

distribution across the landscape on and around agricultural

fields, results in, on average, more than 71% soil loss reduction

(with variations ranging between 50% and 93% in specific con-

texts) (Table 1; Figure S1). The exact value for this contribution is

driven by the mechanics of soil particle detachment, soil

covering vegetation, or litter, which, in theory, should include

coverage across all surfaces.

For particle or nutrient interception by riparian buffers, the

spatial configuration requirements we synthesized are quite spe-

cific, concentrating on the margins of rivers and streams.

Despite variations depending on the levels of pollutants and

sediment, our review indicates that vegetative buffers of at least

28 m, located on both sides of a stream headwater and close to

the water body, notably on slopes <23�, on average, are gener-

ally able to capture more than 73% of non-point-source pollut-

ants (with variations ranging between 50% and 90% depending

on the context) (Table 1; Figure S1). These pollutants include

sediment, nutrients, pesticides, and salts from upstream agricul-

tural lands.

We did not identify and review a specific maximum distance

from habitat for enhancing slope stability and reducing land-

slide occurrence on steep terrains (slopes > 35�) due to the

lack of a straightforward relationship between slope stability

and the distance between habitat and the locations of potential

erosion. Nevertheless, the position and distribution of habitat

are more crucial factors in these landscapes than merely the

distance between the benefiting area and the habitat. Some
64 One Earth 7, 59–71, January 19, 2024
studies indicate that, on these slopes, re-

taining at least 50% complex, diverse

(semi-)natural vegetation cover, distrib-

uted evenly with trees (the heaviest ele-
ments) placed mainly on the base or the bottom of the slope,

is most effective.29,30,35

Functional integrity levels
Based on our review of minimum levels for habitat quantity,

quality, and spatial configuration across the six NCP assessed,

we propose a general integrative measure of functional integrity

that underpins the provisioning of multiple NCP in human-

modified landscapes. We emphasize that our methodology

focuses on the minimum levels of habitat quantity, quality,

and spatial configuration necessary for securing NCP provision

across diverse landscapes. This is distinct from attempting to

quantify the optimal levels needed to meet demand. When

habitat quantity, quality, and spatial configuration are com-

bined, we estimate that at least 20%–25% complex, diverse

(semi-)natural habitat is required for each km2 in human-modi-

fied landscapes to secure ecological functions underlying mul-

tiple NCP provision (Table 1). This estimate is based on the

minimum needs across the six NCP, below which NCP provi-

sion experiences a strong decline (Table 1). Requirements for

individual NCP were determined by the median of the values

reported in individual studies. Using the median implies that,

in certain contexts, a higher quantity of (semi-)natural vegeta-

tion is needed, while in others a smaller area might be sufficient

(see Note S1 for more details). The results further indicate that,

for areas with high erosion or landslide risk, a greater habitat

fraction is necessary (50% habitat per km2). Conversely, in spe-

cific contexts such as doubling crop diversity, or for some spe-

cific NCP such as water quality regulation, NCP provision may

still be achieved with habitat areas as low as 10%–20%. This

variation in the exact quantity required underlines the critical

need to adapt these estimates to align with the specific condi-

tions of each location, accounting for both the local context

and the demand for NCP. However, our analysis indicates

that, below 10% habitat level, NCP provision becomes practi-

cally absent across five NCP, as shown in Figure 1, indicating

that ecosystem functions supporting NCP provision become

unviable below this level. This finding is revealed in 95% of

the studies that we reviewed based on habitat quantity.



Figure 4. Current state of functional integrity in human-modified lands

Habitat functional integrity in human-modified lands (agricultural and urban landscapes) calculated as the percentage (%) of (semi-)natural habitat within 1 km2.

Functional integrity is calculated at a 10-m resolution and then aggregated for display purposes. (A) The global spatial distribution of biosphere functional integrity

at a 500-m scale. More detailed views are shown in the zoom-in panels at a 100-m resolution for (B) East-African highlands and savannah, (C) Argentinian soybean

region, (D) west-central Europe, and (E) Indian Gangetic plain. Areas colored white indicate regions where there are no human-modified lands in our analysis.

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OPEN ACCESSArticle
Current state and spatial distribution
Using the European Space Agency (ESA) 10-m resolution land-

cover map of openly available satellite-based land-cover data,

we estimated the current state and spatial distribution of func-

tional integrity by calculating the percentage of habitat per

1-km2 neighborhoods after distinguishing pastureland from

(semi-)natural grasslands and testing for distinguishing forest

plantations from (semi-)natural forests.36 Our results indicate

that about 50% of human-modified lands are below the level

of 10% habitat per km2, and 64% and 70% of human-modified

lands are below 20% and 25% habitat per km2, respectively

(Figure 4; Table S2). This implies that 20% of human-modified

lands have a habitat area between 10% and 25% per

km2.These areas are likely to have only a limited provision of

NCP, depending on their dependence on external inputs (e.g.,

pesticides) and their vulnerability to climate change. Hence,

only 30% of human-modified lands meet the minimum level

for NCP provision with embedded habitat exceeding 25%

habitat per km2. A significantly higher area than previously esti-

mated, using lower-resolution imagery, has insufficient func-

tional integrity.12 While the limited thematic resolution of the

land-cover data and assumptions made in the analysis may

lead to an underestimation of habitat in the landscape, particu-

larly in terms of small-scale elements (i.e., floral resources,

grassy patches, and hedgerows), it is likely that approximately

two-thirds of all global human-modified landscapes fall below

the 20% per km2 minimum required to provide essential NCP

and are thus heavily reliant on substitutes for those NCP
(domesticated honeybees, pesticides, technical means of wa-

ter regulation and purification) or face absolute shortages in

NCP. This shortage is especially found in the intensively farmed

regions important to global food systems, threatening the long-

term resilience and adaptive capacity of food production

systems.

DISCUSSION

Conservation initiatives have often overlooked the crucial role of

biodiversity in delivering and supporting key NCP that underpin

human well-being, specifically in human-modified lands. Our

study synthesized existing literature to determine the minimum

quantity, quality, and spatial configuration of (semi-)natural

habitat needed in human-modified landscapes for securing the

ecological functions that underlie NCP provision, referred to as

functional integrity. Below these minimum levels, there is a

high risk that human-modified landscapes experience a severe

decline in their capacity to support NCP provision, jeopardizing

the well-being of those dependent on these NCP.

Implications of results
Our findings add to a growing body of evidence, suggesting

that the decline of biodiversity under certain levels is contrib-

uting to a significant decline in NCP provision for the people

who rely on them.15,37 We find that a median of at least 20%–

25% (semi-)natural habitat per km2 (ranging from 6% to 50%

for individual NCP depending on the context) is needed in
One Earth 7, 59–71, January 19, 2024 65



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human-modified landscapes to support multiple NCP provision.

Below a 10% habitat per km2 level, almost all studies indicated

that five of the studied NCP provisions sharply declined to a

very low level or were almost completely absent. For individual

NCP, a lower level sometimes is possible. For example, for wa-

ter quality, 6% might be sufficient, based on the minimum buffer

width required for riparian strips, depending on the drainage

density of the area and steepness.

NCP are delivered by communities of species across taxa and

their traits. Vegetation characteristics define habitat quality and

provide biophysical contributions such as sediment interception

while providing resources for mobile species that contribute to

pollination and pest control. Our analysis aligns and extends

the existing knowledge by focusing on the incorporation of com-

plex diverse (semi-)natural elements (Table 1; Table S1).

Although many NCP can be procured with non-native spe-

cies,38,39 incorporating embedded habitats that promote native

species, and improving connectivity within fragmented land-

scapes can provide additional biodiversity conservation bene-

fits.40 This also supports the protection of cultural heritage and

local knowledge.41 Complementary to this, increasing intra-field

diversity of the agricultural/modified elements26,42 and field edge

density22 and decreasing field sizes26 may also increase land-

scape heterogeneity but does not replace the positive effect of

the area of (semi-)natural elements on functional integrity. Which

practices, types of habitats, or landscape elements are most

appropriate to ensure functional integrity remains a highly local

issue and requires input from local knowledge.43

Ensuring access to appropriate habitat (or landscape ele-

ments) at a sub-kilometer scale is important across all human-

modified landscapes. Larger areas in one place cannot substi-

tute for smaller areas in another. This is driven by the fact that

the majority of species providing NCP have small home ranges

or are non-mobile. Numerous ecological studies also show

non-linear decreases in species diversity and abundance with

increasing distance from habitat edges.44,45 An additional

benefit of embedding habitats within human-modified land-

scapes is the fragmentation of large areas of agricultural lands.

This strategy reduces the dispersal of agricultural pests between

fields46 while connecting habitat of species that can reduce pest

pressure.47 It also contributes to reducing soil erosion and

improving soil biological activity and fertility. Securing riparian

buffers is a good first step and would, for example, secure about

6% of habitat per km2 on average globally, while contributing to

connectivity.48

The minimum functional integrity level identified here is appli-

cable tomost human-modified landscapes that have demand for

one or more of the considered NCP.15,28,37,49 Local demand for

specific NCP can vary strongly, depending on factors such as

cropping systems’ dependence on pollination, topography, pop-

ulation density, and societal needs.50–52 Rather than identifying

the required supply to meet this demand, our study identifies

the minimum level of functional integrity necessary to secure

ecological functions in human-modified lands, emphasizing bio-

diversity’s functional contributions in supporting both regulating

and non-material NCP at local scales. This includes contribu-

tions that either improve food production or reduce its negative

environmental impacts, as well as those that promote people’s

mental and physical well-being. Meeting NCP demand, in
66 One Earth 7, 59–71, January 19, 2024
many conditions and contexts, will require habitat quantity, qual-

ity, and spatial distribution levels that are greater than the mini-

mum values identified here. Therefore, local implementation

needs to go beyond this analysis and adjust the requirements

for diverse socioecological contexts and specific NCP demands

and relations between NCP supply and ecological functions.

Engaging with local communities and implementing locally

adapted practices are fundamental steps in identifying which

NCP to prioritize and the critical habitats that provide them to

ensure effective conservation strategies and foster equitable

and sustainable ecological practices.43

Methodological considerations
Although our review approach may have overlooked some

important primary research articles, this is unlikely to have influ-

enced our results, as the results indicate a high level of agree-

ment among the current, large, body of evidence. The analyzed

studies cover a diverse range of locations across the globe.

Nevertheless, these types of studies often reveal strong biases

due to the locations where primary research is conducted as is

common in ecological research, suggesting that some biomes

might be underrepresented.53 While we did not fully capture all

facets of biodiversity and NCP that are essential for supporting

human needs, the majority of the NCP we selected represent a

core set of regulating ecosystem functions that are important

at local scales and essential for human well-being. However,

functional integrity as operationalized in this study is unable to

capture finer-scale NCP provision, notably those related to soil

biodiversity and ecosystems. These include soil quality, below-

ground carbon sequestration, nutrient cycling, and increased

water-holding capacity in fields. This is evident in the higher min-

imum level identified for soil and canopy cover to prevent soil

particle detachment (>50% vegetation cover). Our measure of

functional integrity does not also capture complementary prac-

tices that can either improve NCP production or reduce pres-

sures on habitat to provide NCP. For example, no-till or

reduced-tillage practices, improved nutrient-use efficiency,

cover crops, or leguminous rotations reduce erosion and nutrient

loss but are not captured by the metric we proposed. Comple-

mentary metrics and practices incorporating soil biodiversity

and soil-based NCP are equally important and call for greater

integration of ecological principles across all land surfaces.54–56

While field-scale practices that reduce excess nutrient run-off

directly from human-modified lands (e.g., field tillage practices)

are important, they complement but do not replace the role of

habitat in buffering soil, nutrient, and pollutants’ loss to aquatic

ecosystems.57 It is also important to note that excessive nutrient

application can rapidly exceed the absorption capacity of ripar-

ian and other vegetated buffers; therefore, reducing such pres-

sures can increase the capacity of habitat to maintain functional

integrity.

Historically, global monitoring of functional integrity of human-

modified landscapes has been challenging, as habitat mostly

comes in small patches, often of linear format, that are not easily

detectable in most coarse-resolution global (and regional) land-

cover maps. The recent high-resolution Sentinel images (10-m

resolution) used here can capture relatively small patches. How-

ever, even these datamight still underestimate habitat as they do

not capture linear elements such as hedgerows, field margins,



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OPEN ACCESSArticle
floral strips, and grass strips that are managed as (semi-)natural

habitat. This is partially due to the limited spatial resolution but

also a result of the limited thematic resolution of this data prod-

uct and the absence of information on vertical or 3D structures

(e.g., vegetation height). For example, unmanaged patches of

grassland are not sufficiently distinguished from pasture and

(semi-)natural grasslands, and low-intensity pastureland may

have good ecological condition. Similar concerns hold for forest

land cover through remaining challenges of distinguishing be-

tween different types of forests. For example, we could not

distinguish natural forests from monocultures of short-rotation

species in our analysis. We tested the sensitivity of the results

to distinguishing forest plantations from other forests. While

the impact on the global results was small, it did show clear

regional deviations (Table S2). Finally, the results of our analysis

are sensitive to the classification of bare lands as either being a

natural habitat or a human-modified land cover, which is not

distinguished in the data used. Given these limitations, our

assessment of the current state of functional integrity should

be interpreted with caution. We anticipate that, with continued

rapid evolution of remote-sensing products and artificial intelli-

gence, these detection challenges will be reduced in the near

future. Early analysis of satellite imagery using deep learning to

monitor highly heterogeneous areas have been published58 but

are not yet openly available for inclusion in our assessment.

Current state and habitat restoration pros and cons
Despite the challenges of detecting small-scale landscape ele-

ments in highly heterogeneous areas, as highlighted by the

aforementioned issues, we estimate that at least two-thirds of

the global human-modified lands fall below critical levels for

functional integrity, severely compromising the capacity of hu-

man-modified lands to contribute to NCP provision. In agricul-

tural and urban landscapes, the natural vegetation has frequently

been removed to accommodate the growing demands for hous-

ing and agricultural production. Competition for land may limit

space for restoring natural elements. Therefore, restoring habi-

tats in these places is often interpreted as conflicting with the

provision of material NCP and might compete with ambitions

of increasing food production as well as with the needs and pri-

orities of local communities (e.g., housing). In reality, this

perceived conflict does not always preclude mutually beneficial

outcomes, and the magnitude and direction of the effect can

vary largely depending on local context. The literature shows ev-

idence that, in many places, a diversity of practices improve both

yields and environmental outcomes and that embedded biodi-

versity on field perimeters and riparian buffers leaves scope for

sustainable intensification within fields.59–61 Well-functioning

ecosystems can support the provisioning of material NCP

through contributions such as climate regulation, nutrient

cycling, and pest control. These contributions are particularly

useful when reducing the environmental impact of pesticides

and fertilizers is necessary. Therefore, the generalized trade-off

between the area of natural habitat and food production is, in

some cases, a misconception. Locally appropriate conservation

options can be effective in managing and mitigating potential

conflicts between material and non-material NCP. Implementing

notably modern agroecological practices and nature-based so-

lutions can help to better integrate new habitats in these land-
scapes and minimize trade-offs. Adopting diverse crop rotations

and mixed cropping systems maintains habitat heterogeneity,

supports various species, and promotes ecosystem resil-

ience.62,63 Other benefits of habitat in human-modified land-

scapes include the significant contributions of an increase in

tree cover in agricultural landscapes (e.g., agroforestry systems)

to soil health, water retention, and global carbon sequestra-

tion.64 Strategically placing small patches of habitat in human-

modified landscapes, combined with innovative techniques

such as precision agriculture practices, may also have dispro-

portionate value in preserving species diversity65 while also opti-

mizing agricultural productivity.66

Conclusions and future directions
Restoring habitats and their ecosystem functions in human-

modified landscapes can help strengthen the resilience of these

areas toward climate change.67 Therefore, the benefits of at least

meeting the minimum habitat level identified in our study offer

broader benefits that extend beyond the analyzed NCP. Notably,

our critical finding (supported by 95% of the studies we re-

viewed) that NCP provision is likely to be largely absent for five

NCP when habitat level drops below 10% of the landscape em-

phasizes the urgent need for policy intervention in areas with

habitat below that level. We stress that the minimum level iden-

tified here is a minimal requirement to secure ecosystem func-

tions underlying multiple NCP provision, rather than an optimal

level required to meet demands for NCP. Contextualized strate-

gies, responding to local demands for NCP, can further optimize

the benefits of such habitat within the landscape and contribute

to safeguarding biodiversity, promoting ecosystem stability, and

contributing to overall human well-being. The shortcomings

of (semi-)natural habitat in many landscapes across the globe

reconfirm the high importance of not only focusing conservation

and restoration efforts on intact natural or wilderness areas.

Conservation and restoration efforts, especially in the UN

decade of restoration, also have prime importance in strongly

modified landscapes.

EXPERIMENTAL PROCEDURES

Resource availability

Lead contact

Further information and requests for resources should be directed to and

will be fulfilled by the lead contact, Awaz Mohamed (awaz.mohamed2@

gmail.com)

Materials availability

This study did not generate new unique materials.

Data and code availability

All datasets and codes generated in this study to estimate the current

state and to produce the maps, as well as the full list of studies used in

NCP analysis, have been deposited at DataversNL under https://doi.

org/10.34894/V6WWTS, and are publicly available as of the date of pub-

lication. The dataset generated to estimate functional integrity level and

to produce the figures will be shared openly by the lead contact upon

request after publication.

Methods

NCP selection

We selected NCP for human-modified lands that are underpinned by various

specific ecological processes. Notably, we focused on five regulating and

one non-material NCP that are related to biodiversity and ecosystem function

and directly affect the well-being of local people and their quality of life in
One Earth 7, 59–71, January 19, 2024 67

mailto:awaz.mohamed2@gmail.com
mailto:awaz.mohamed2@gmail.com
https://doi.org/10.34894/V6WWTS
https://doi.org/10.34894/V6WWTS


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OPEN ACCESS Article
different manners, from regulating the quality of water, to pollinating many

crops, to underpinning multiple dimensions of human health. These include

(1) pollination; (2) pest and disease control; (3) physical and psychological ex-

periences in nature, termed ‘‘experiences 2; (4) soil erosion control; (5) water

quality regulation; and (6) natural hazards mitigation. We define human-modi-

fied lands as the inhabited, used, and working lands of the world (e.g., heavily

modified anthromes) where the ecosystem is dominated by human activities

that have largely changed the natural ecosystem functions and composition.

In our study, we considered a wide range of human-modified lands, including

urban areas, forest plantations, and agricultural lands.

Literature search strategy

We conducted a literature search of peer-reviewed reviews and meta-ana-

lyses following the Preferred Reporting Items for Systematic Review and

Meta-Analyses guidelines (PRISMA).68 We employed two to three keywords

and terms standardized, combining specific NCP, habitat or vegetation, and

landscape scale. We analyzed (1) pollination (‘‘pollinat*’’ AND ‘‘habitat’’ AND

‘‘landscape), (2) pest and disease control (‘‘Biological control*’’ AND ‘‘habitat’’

AND ‘‘landscape’’), (3) physical and psychological experiences (‘‘physical AND

psychological*’’ AND ‘‘well-being*’’ AND ‘‘nature*’’), (4) water quality regulation

("riparian buffer*" AND "width*"), (5) soil erosion control ("soil erosion*" AND

"vegetation*" AND "landscape*"), and (6) natural hazards ("landslide*" AND

"vegetation cover*"). All searches were conducted on Web of Science (Clari-

vate Analytics, Philadelphia, PA, USA). Additional reviews and primary papers

were identified from other sources, whether suggested by experts (regardless

the year of publication) or similar searches on Google Scholar engine using an

additional search string for each NCP (e.g., ‘‘pollination OR pollinators*’’;

habitat*; landscape configuration*; landscape complexity*; landscape hetero-

geneity*). We screened the first two pages of Google search results, selecting

relevant articles based on the titles and abstracts (Figure S2). The output of

these queries was saved to the Zotero open-source software (Zotero site:

www.zotero.org), where all papers and citations were managed.

Eligibility criteria

Before proceeding with the evaluation process, we established predetermined

criteria for inclusion. We adhered to the following guidelines to select poten-

tially relevant references for subsequent stages of evaluation: Reviews and

meta-analyses needed to be published in peer-reviewed journals between

January 2010 and December 2021 and be in English. Each source should

focus on which taxonomic groups provide that NCP, the area and the quality

of (semi-)natural habitat including relevant landscape elements, the distance,

location, or placement of landscape elements, and a description of the spatial

relationship between biodiversity and the specific NCP. We assumed that the

majority of review papers published between 2010 and 2021 were built on pri-

mary research articles, some of these originating from before 2010. There were

no restrictions imposed concerning methods used, landscape type, or study

location. Any references not fulfilling any of the above criteria or clearly being

out of scope (i.e., did not report outcomes on habitat quantity, quality, or

spatial configuration) were excluded from the analysis. However, for some

NCP where only a limited number of review papers was available (%5 reviews,

covering <50 original papers in total), such as experiences or landslide mitiga-

tion, we randomly incorporated a selected number of primary research articles

from experts and other sources to ensure a better balance in the number of pa-

pers included across different NCP. Our preliminary search yielded 411 papers

in total after duplicates were removed.

Study selection

We initially screened all papers (n = 411) based on titles and abstracts to iden-

tify potentially eligible and relevant reviews. Abstracts that did not fulfill at least

one of our aforementioned inclusion criteria or were deemed irrelevant to the

topic upon closer inspection were excluded. We then skimmed through the

full-text articles to further assess the quality and relevance. We included pa-

pers addressing at least one of three key variables for each NCP to delineate

the minimum level of functional integrity that secures the ecosystem functions

underlying the NCP. These encompass (1) a quantitative measure of the min-

imumhabitat required for supporting NCP provision, (2) a qualitative evaluation

of landscape elements’ type and quality required, and (3) the maximum dis-

tance between providers and beneficiaries (in meters) or the spatial configura-

tion of landscape elements required for supporting NCP provision. Papers that

did not meet the inclusion criteria were excluded (Figure S2). Our search

yielded a total of 154 articles (the full list of publications is available in the re-
68 One Earth 7, 59–71, January 19, 2024
pository, under the ‘‘data and code availability’’ section), comprising 74

meta-analyses and reviews and 80 primary research articles conducted in

different locations around the world (Figure S2). While acknowledging that

some primary articles may have been included in multiple reviews and meta-

analyses, we could not verify this across all reviews. To further reduce biases

due to unequal levels of evidence in review papers, we weighted the calcula-

tions by the number of papers included in each review andmeta-analysis used.

Data extraction and management

We extracted data from all eligible reviews and original articles and tabulated

them using a set of data extraction forms developed for this study. We gath-

ered the following information: name of the first author, publication year, jour-

nal name, study’s location, nature of the paper, number of papers included,

estimated minimum habitat quantity, description of habitat elements, land-

scape elements recommended, estimated maximum linear distance from

source habitat or the location and emplacement of habitat, functional group

providing the NCP, slope, buffer or vegetation cover efficiency in reducing

soil loss or non-point pollutants, and estimated minimum buffer width.

Minimum level estimation

In the reviewed studies, we determined the minimum habitat value in the land-

scape below which the function underlying each NCP shows a strong decline

or is almost completely absent. This determination was based on the informa-

tion reported in the text, tables, figures, and supplemental information pro-

vided in the individual papers. Below this minimum habitat level, certain

ecological functions may lack resilience, with the habitat possibly being insuf-

ficient to maintain ecological functions due to the loss of critical species or

viable population levels to overcome shocks. For pollination and pest and dis-

ease control NCP, when a figure displayed the relationship between the abun-

dance or diversity of NCP-providing organisms and habitat area, we deter-

mined the minimum area of habitat quantity by identifying the point where

their abundance or diversity strongly declined to zero or a minimal value

slightly above zero (Figure S3). This decline in the abundance and diversity

of NCP-providing organisms indicates a strong decline to very low level or

the complete loss of the ecological functions or associated NCP. For benefits

from experiences, we assessed the minimum amount of green spaces of

various forms and qualities required in urban ecosystems, considering their

spatial configuration or linear distance (see Table 1) from each neighborhood.

These values were derived from studies examining the link between the

amount of green space in each neighborhood in cities and people’s mental

and physical well-being. These studies measured variables such as psycho-

logical distress level, number of natural-cause mortality, cortisol levels, pre-

scriptions for antidepressants, presence of anxiety, COVID-19 incidence

rate, and heat stress level69–71 as indicators of mental and physical well-being.

We determined the minimum area of habitat quantity by pinpointing the point

where these aforementioned variables sharply dropped to zero or nearly zero.

In general, habitat quantity estimation was made irrespective of variations in

contexts, methods, relationship types, or locations between studies.

For soil erosion control and water quality regulation, we examined studies

exploring the efficiency (measured as percentage) of vegetation cover and

vegetated buffers in reducing soil loss and pollutants. However, efficiency

varies highly between studies and depends on the specific NCP and land-

scape type. Hence, there is no universally agreed-upon minimum efficiency

level proposed across these studies. In 90% of the reviewed studies, different

amounts of vegetated buffers exhibited efficiency exceeding 50%. Therefore,

we adopted >50%efficiency as a baseline in our analysis to determine themin-

imum required vegetation value. The buffer width was represented in meters.

To transform this buffer width into an approximate amount of (semi-)natural

vegetation per km2, comparable with other NCP, we used the global average

density of streams.72

For natural hazard mitigation, particularly landslide mitigation, the minimum

value level of habitat quantity required has been derived from experimental

and modeling studies assessing the factor of safety (FoS) in relationship to

the presence and absence of plant roots in the soil.33,73–75 FoS is a crucial indi-

cator of slope stability and is defined as the ratio of the resisting force to the

driving force along a failure surface.75 Maintaining a slope stability often re-

quires a FoS value of 1.3 for temporary or low-risk slopes and 1.5 for high-

risk slopes.76 Therefore, we use the 1.3 FoS as a baseline in our analysis to

determine the minimum vegetation cover required for maintaining slope

stability.

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OPEN ACCESSArticle
To assess habitat quality, we analyzed the literature collected that recom-

mended various landscape elements essential for the survival of individuals

and the persistence of populations that contribute to these NCP. Based on

our analysis, we identified common landscape elements across the reviewed

studies and classified them into six categories, guided by each paper’s recom-

mendation and different contexts. These are complex diverse (semi-)natural

habitat, complex diverse natural habitat, diverse floral resources, forest,

grassy elements, and woody elements (Table S1 for more details). Some

studies broadly described these categories without mentioning any specific

landscape elements, while others provided more specific descriptions,

mentioning particular landscape elements.

To assess spatial configuration needs for three of the six NCP provided by

mobile organisms (pollination, pest and disease control, and experiences),

we estimated the maximum linear distance these mobile organisms forage

or travel from their home habitat to access resources. For the remaining three

NCP provided by non-mobile organisms (water quality regulation, soil erosion

control, and natural hazard regulation), we extracted recommended habitat

placement and location descriptions that support NCP.

Once minimum habitat quantity, quality, and spatial configuration values

were established for each NCP at the landscape scale, we performed explor-

atory analyses to identify general patterns in the literature regarding the three

key variables for each specific NCP. All analyseswere conducted using the Py-

thon language (Python 3.6) within the Anaconda platform (Seaborn and mat-

plotlib libraries). We then assessed which characteristics of functional integrity

(habitat quantity, habitat quality, and spatial configuration essential for func-

tioning) are important for decision makers and management.

Functional integrity: Current state and spatial distribution

We assessed the current state of the functional integrity level using the ESA

WorldCover 10-m resolution land-cover map (https://esa-worldcover.org/

en). First, we created a binary map of what entails habitat within human-modi-

fied landscapes. We refined the grassland category to distinguish pasture-

lands from (semi-)natural grasslands by overlaying the habitat map from

Jung et al.36 Specifically, areas classified as grassland in the ESA

WorldCover 10-m resolution land-cover map, which overlapped with the

area classified as ‘‘artificial – terrestrial’’ by Jung et al.,36 were reclassified as

pastureland. All remaining grassland areas were reclassified as ‘‘natural grass-

land.’’ We then reclassified the refined map to create a binary classification of

‘‘natural lands’’ and ‘‘human-modified lands’’ (Table S3). As this is based on

land-cover data only, we acknowledge the likely underestimation of human

modification of nature in this map. We calculated a functional integrity value

for each pixel using a focal function where we calculated the percentage of

habitat cover in a 500-m radius around each pixel. We calculated the percent-

age of pixels that met or exceeded different critical levels of functional integrity

(10%, 20%, 25%) on a global scale and on an ecoregion scale. Furthermore,

we performed an additional sensitivity analysis using the Jung et al.36 classifi-

cation to refine the ESA WorldCover 10-m resolution land-cover map ‘‘tree

cover’’ category. We reclassified pixels where the ESA WorldCover 10-m res-

olution land-cover area classified as tree cover overlapped with areas classi-

fied as plantations by Jung et al.36 All other ESA WorldCover 10-m resolution

land-cover tree-cover pixels were reclassified as natural tree cover. Natural

tree cover was assigned a 1 and ‘‘plantations’’ was assigned a 0 in the binary

classification. We then followed the same procedure as above to calculate

functional integrity value (Table S2). All analyses were done using the native

Google Earth Engine interface.

SUPPLEMENTAL INFORMATION

Supplemental information can be found online at https://doi.org/10.1016/j.

oneear.2023.12.008.

ACKNOWLEDGMENTS

This work is part of the Earth Commission, which is hosted by Future Earth and

is the science component of the Global Commons Alliance. The Global Com-

mons Alliance is a sponsored project of Rockefeller Philanthropy Advisors,

with support from Oak Foundation, MAVA, Porticus, Gordon and Betty Moore

Foundation, Tiina and Antti Herlin Foundation, William and Flora Hewlett Foun-

dation, Generation Foundation, and the Global Environment Facility. The Earth
Commission is also supported by the Global Challenges Foundation and Fron-

tiers Research Foundation. Individual researchers were supported by the Eu-

ropean Research Council grant on Climate Change and Fossil Fuel (project

number 101020082) (J.G.) and the Open Society Foundations (J.F.A.). F.D.C.

received additional support from the Food System Economics Commission

and PHV from the NatureConnect project funded by the European Commis-

sion Horizon Europe program. F.D.C., S.J., and N.E.C. acknowledge funding

support from the OneCGIAR NEXUS Gains initiative, Realizing Multiple Bene-

fits AcrossWater, Energy, Food and Ecosystems (Forests, Biodiversity). I.C.M.

acknowledges the Deutsche Forschungsgemeinschaft (DFG, German

Research Foundation) financial support awarded within the Heisenberg pro-

gram (grant no. ME 4156/5-1). We also acknowledge Nuno Martins Migueis

Garcia for his invaluable assistance with the spatial analysis and data

repository.
AUTHOR CONTRIBUTIONS

A.M. developed themethodology for assessing and analyzing functional integ-

rity, participated in the conceptual design, conducted the systematic review,

gathered and analyzed data, led the write-up of the paper, and served as a

research scientist on the Earth Commission’s Biosphere interactions working

group. F.D.C., P.H.V., and D.O. originated the idea, developed the concept

and methodology for assessing functional integrity, contributed to the analysis

and write-up, and co-led the Earth Commissions Biosphere Working Group.

J.F.A. participated in the conceptual design and writing of the paper, per-

formed the spatial integrity analysis, created the spatial maps, and served

on the Earth Commissions Biosphere Working Group. N.Z.C. contributed to

the analysis and the writing of physically and psychologically beneficial expe-

riences in nature. N.E.-C., A.F., and S.J. contributed to the conceptualization

andmethodology for assessing functional integrity and to reviewing of the final

manuscript. J.R. participated in the conceptual design and writing of the paper

and served on the Earth Commissions Biosphere Working Group. I.C.M.

contributed to the analysis of the soil erosion control NCP and to reviewing

the final manuscript. B.S.K. contributed to the riparian analysis and reviewing

the final manuscript.
DECLARATION OF INTERESTS

The authors declare no competing interests.

Received: December 6, 2022

Revised: April 17, 2023

Accepted: December 7, 2023

Published: January 19, 2024

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