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Ten golden rules for reforestation to optimize carbon sequestration, biodiversity recovery and livelihood benefits



Urgent solutions to global climate change are needed. Ambitious tree‐planting initiatives, many already underway, aim to sequester enormous quantities of carbon to partly compensate for anthropogenic CO2 emissions, which are a major cause of rising global temperatures. However, tree planting that is poorly planned and executed could actually increase CO2 emissions and have long‐term, deleterious impacts on biodiversity, landscapes and livelihoods. Here, we highlight the main environmental risks of large‐scale tree planting and propose 10 golden rules, based on some of the most recent ecological research, to implement forest ecosystem restoration that maximizes rates of both carbon sequestration and biodiversity recovery while improving livelihoods. These are as follows: (1) Protect existing forest first; (2) Work together (involving all stakeholders); (3) Aim to maximize biodiversity recovery to meet multiple goals; (4) Select appropriate areas for restoration; (5) Use natural regeneration wherever possible; (6) Select species to maximize biodiversity; (7) Use resilient plant material (with appropriate genetic variability and provenance); (8) Plan ahead for infrastructure, capacity and seed supply; (9) Learn by doing (using an adaptive management approach); and (10) Make it pay (ensuring the economic sustainability of the project). We focus on the design of long‐term strategies to tackle the climate and biodiversity crises and support livelihood needs. We emphasize the role of local communities as sources of indigenous knowledge, and the benefits they could derive from successful reforestation that restores ecosystem functioning and delivers a diverse range of forest products and services. While there is no simple and universal recipe for forest restoration, it is crucial to build upon the currently growing public and private interest in this topic, to ensure interventions provide effective, long‐term carbon sinks and maximize benefits for biodiversity and people.
Glob Change Biol. 2021;00:1–21.
Received: 25 Augus t 2020 
Accepted: 13 Octob er 2020
DOI: 10.1111/gcb.15498
Ten golden rules for reforestation to optimize carbon
sequestration, biodiversity recovery and livelihood benefits
Alice Di Sacco1| Kate A. Hardwick1| David Blakesley2,3| Pedro H. S. Brancalion4|
Elinor Breman1| Loic Cecilio Rebola1,5 | Susan Chomba6| Kingsley Dixon7, 8 |
Stephen Elliott9| Godfrey Ruyonga10| Kirsty Shaw11| Paul Smith11 |
Rhian J. Smith1| Alexandre Antonelli1,12,13
1Royal Botanic Gardens, Kew, Richmond, UK
2Wildlife Landscape s, Maidstone, U K
3Autism and Nature, Maidstone, UK
4Depar tment of Fore st Scien ces, “Luiz de Queiroz” College of Agriculture, Unive rsity of São Paulo, Piracicaba, SP, Brazil
5School of Biological Sciences, University of A berdeen, Aberdeen , UK
6World Agrofores try Centre, N airobi , Kenya
7Austral ian Rese arch Council Centre for Mi ne Site Restorat ion, School of Mo lecula r and Life Sciences, Cur tin Uni versit y, Perth , WA, Aus tralia
8Missouri Botanical Garden, St Louis, M O, USA
9Forest Re storation Research Un it and Environmental Science Res earch Centre, Biolog y Depar tment, Facult y of Scie nce, Chiang Mai University, Chiang M ai,
10Tooro Botanical Gardens , Fort Po rtal, Ugand a
11Botanic Garde ns Conservat ion International, Richmond , UK
12Depar tment of Biologi cal and Environmental Science s, Gothenbur g Globa l Biodiversit y Centr e, University of Gothenburg, Gothenburg, Sweden
13Depar tment of Plant Scie nces, University of Ox ford, O xford, UK
This is an op en access arti cle under the ter ms of the Creative Commons Attribution L icense, which pe rmits use, dis tribu tion and reprod uction in any med ium,
provide d the original wor k is properly cited.
© 2021 Royal Botanic G arden, Kew. Global Change Biology published by John Wiley & Sons Lt d
Alice Di S acco and Kate A. H ardwick shou ld be cons idered joint fi rst author.
Kate A. Hardwick and Alexandr e Antone lli,
Royal Bot anic Gardens, Kew, Richmond,
London T W9 3AE, UK.
Email: k. (K. A. H.); (A . A.)
Funding information
Royal Botanic Gardens, Kew; Fundaç ão
de Amparo à Pesquisa do Es tado de
São Paul o, Grant/Award Number:
2018/18416-2; Sky Zero; Swe dish
Research Council, Gra nt/Award Number:
2019-05191; Chiang Mai University;
Swedish Foundation for Strategic
Research, Gra nt/Award Number: FFL15-
Urgent solutions to global climate change are needed. Ambitious tree-planting ini-
tiatives, many already underway, aim to sequester enormous quantities of carbon to
partly compensate for anthropogenic CO2 emissions, which are a major cause of ris-
ing global temperatures. However, tree planting that is poorly planned and executed
could actually increase CO2 e mis sio ns and have long-ter m , deleter iou s impa c t s on bi o-
diversity, landscapes and livelihoods. Here, we highlight the main environmental risks
of large-scale tree planting and propose 10 golden rules, based on some of the most
recent ecological research, to implement forest ecosystem restoration that maximizes
rates of both carbon sequestration and biodiversity recovery while improving liveli-
hoods. These are as follows: (1) Protect existing forest first; (2) Work together (involv-
ing all stakeholders); (3) Aim to maximize biodiversity recovery to meet multiple goals;
(4) Select appropriate areas for restoration; (5) Use natural regeneration wherever
possible; (6) Select species to maximize biodiversity; (7) Use resilient plant material
   DI SACCO et Al .
Trees, and the forests they form, are highly complex. Their interac-
tions with other plants, animals and fungi, and environmental phe-
nomena such as fires and flooding, have led to the evolution of a
remarkable diversity of species, genes, functions and ecosystems.
In Amazonia alone, it has been estimated that there are more than
15,00 0 tree species (ter Steege et al., 2020). Today, trees and forests
provide people with invaluable products and services (Díaz et al.,
2018), including food, medicine, building materials, fibre, shade, rec-
reational space, pollution filtration and flood risk reduction, and they
are essential reservoirs of carbon, water and nutrients.
The escalating and interconnected threats of biodiversity loss
through deforestation, global climate change (GCC) and pover ty
have increased awareness of the mitigating role that forests could
play (Brancalion & Holl, 2020) and have led to some notable global
initiatives. (Key terms are highlighted in bold on their first occurrence
and defined in Table 1.) The role of forest restoration in GCC mitiga-
tion first received global recognition in 20 08, when ‘enhancement of
forest carbon stocks’ was added to the United Nation's REDD+ ini-
tiative (UNFCCC, 2008;, with measures to ensure
biodiversity conservation and community participation (UNFCCC,
2011; safeguards [d] and [e]). In 2011, the Bonn Challenge (www.
bonnc halle was launched, aiming to restore 350 million ha of
forest globally by 2030. Currently, more than 70 pledgers from more
than 60 countries are restoring 210 million hectares of degraded and
deforested lands ( In 2020, the
World Economic Forum instigated an ambitious global tree-plant-
ing programme—the platform—to support the UN Decade on
Ecosystem Restoration 2021–30 (www.decad eonre stora
These initiatives mostly advocate forest (and) landscape restoration
(FLR)—an approach that aims to ‘regain ecological functionalit y and en-
ha n ce hu man well -bei n g in defo r e ste d or de g rade d lan dsc a pes’ (Be sse a u
et al., 2018). However, concerns are growing that several ambitious
initiatives are falling short of delivering on the three key objec tives of
carbon sequestration, biodiversity recovery and sustainable livelihoods
(e.g. Figure 1; Lewis et al., 2019). They may have set unrealistically high
targets (Fagan et al., 2020) and may have unforeseen negative conse-
quences. Potential problems include displacement of native biodiversity,
particularly due to the destruction of non-forest ecosystems (Seddon
et al., 2019); increases in invasive species (Kull et al., 2019); a reduction
in pollinator services (Ricketts et al., 2004); a reduction in croplands and
thus food production; disruption of water cycles; a decrease in c arbon
stored in aboveground biomass (Heilmayr et al., 2020); a reduction in
soil organic carbon (SOC; Hong et al., 2020; Veldman et al., 2019) and
a lowering of albedo in boreal zones, causing temperature rises (Bett s,
2000). These negative outcomes are mostly associated with the ex-
tensive use of exotic monoculture plantations, rather than restoration
approaches that encourage a diverse, carbon-rich mix of native tree
spe cie s (Br ancalion et al. , 2018; Heilmay r et al., 202 0; Lewis et al., 2019).
Lewis et al. (2019) estimated that only a third of commitment s und er th e
Bonn Challenge and other schemes aim to restore natural forests.
In naturally forested regions that have become deforested by
human activities, we propose a ‘native forest approach’ to FLR, to
increase carbon sequestration and other ecosystem services, ac-
celerate biodiversity recover y and generate sustainable livelihoods.
This approach emphasizes protec ting and restoring native fores t el-
ements within a mosaic of land uses, which would typically include:
(i) Existing native forest, prioritized for protection, to safeguard
carbon stocks, reduce emissions and conser ve biodiversity;
(ii) Restored native forest, to maximize rates of carbon sequestra-
tion and recovery of biodiversity and ecosystem services, deliv-
ering sustainable economic benefits;
(iii) Livelihood native forest, to maximize economic benefits to local
communities while significantly increasing carbon sequestration,
(with appropriate genetic variability and provenance); (8) Plan ahead for infrastruc-
ture, capacity and seed supply; (9) Learn by doing (using an adaptive management
approach); and (10) Make it pay (ensuring the economic sustainability of the project).
We focus on the design of long-term strategies to tackle the climate and biodiversity
crises and support livelihood needs. We emphasize the role of local communities as
sources of indigenous knowledge, and the benefits they could derive from success-
ful reforestation that restores ecosystem functioning and delivers a diverse range of
forest products and services. While there is no simple and universal recipe for forest
restoration, it is crucial to build upon the currently growing public and private interest
in this topic, to ensure interventions provide effective, long-term carbon sinks and
maximize benefits for biodiversity and people.
afforestation, climate-change mitigation, ecological restoration, forest landscape restoration,
large-scale tree planting, natural regeneration, nature-based solutions, stakeholder
DI SACCO et Al .
TABLE 1 Glossary (terms highlighted in bold on first occurrence in the text)
Ter m Definition
Adaptive management An intentional approach to making decisions and adjustment s in resp onse to new information and changes in
Afforestation Creation of forest on areas not natur ally forested in recent times
Agroforestry Restoration and sustainable management of existing agricultural land through integration of trees in the agricultural
Applied nucleation Planting trees in small groups or ‘nuclei’ and reliance on seed-dispers al out from such nuclei to res tore forest cover
across the entire restoration site
Assisted (or accelerated)
natural regeneration
Managing the process of natural forest regeneration to achieve forest ecosystem recover y more quickly, through
interventions such as fencing, weeding and enrichment plantings
The variabilit y within and between ecosystems, species and genetic material
Composite provenancing The use of a mix of mainly local provenance material with a small amount from dist ant but ecogeographically
matched provenances
Deforestation Destruction and degr adation of forest
Existing native forest Old- and second-growth, degraded and planted forests
Forest (and) landscape
restoration (FLR)
Ongoing process of regaining ecological functionality and enhancing human well-being across deforested or
degraded forest landscapes
Forest restoration Restoration of degraded, damaged or destroyed forested areas (see Restoration)
Framework species
Planting a mix of tree species, typical of the target forest ecosystem, that catalyse forest regeneration by shading
out herbaceous we eds and attracting seed-dispersing animals.
Livelihood native forest Mixed species forest with entirely or mostly native species, managed sustainably to provide local economic benefits
Natural regener ation (NR) The process of natural forest regrow th, which can occur spontaneously following land abandonment or be assis ted
by human interventions (see Assisted Natur al Regeneration)
Nature-based solutions
Actions that involve 'working with and enhancing nature to help address societal goals' (Seddon et al., 2019)
Non-timber forest
produc ts (NTFPs)
Commodities obtained from a forest without logging, for example, fruit, honey, mushrooms , medicinal plants
Old-grow th fore st Also c alled primar y or virgin forest. Forest that has not been recently disturbed
Orthodox seeds Seeds that tolerate drying to 5% moisture content and freezing at −20°C (approximately 92% of all plant species), as
opposed to recalcitrant seeds that do not survive such conditions and would require cr yopreservation (storage
at around −196°C in liquid nitrogen) or direct cultivation
Outcrossing species Species that reproduce by fertilization between gametes produced by dif ferent individuals
Payment s for ecosystem
services (PES)
Financial incentives for managing land that provides an ecological service, for example, watershed protection
Predictive provenancing
(also called provenance
The use of distant genotypes that are experimentally determined to be adapted to projected conditions
Proforestation Protecting existing natural forest s
REDD+ Programme from the United Nations for ‘Reducing Emissions from Deforestation and forest Degradation and the
role of conservation, sustainable management of forest s and enhancement of forest carbon stock s in developing
Reforestation Re-creation of forest on a previously forested area
Restoration ‘The process of assisting the recover y of an ecosystem t hat has been degraded, damaged, or destroyed’ (Gann et al.,
Restored native forest Native forest ecosystems reinst ated on degraded land
Second-growth (or
secondary) forest
Forest grown after recent disturbance
Seed zone An area within which plant materials c an be transferred with little risk of being poorly adapted to their new location
Selfing species Species that reproduce by fertilization between gametes within the same hermaphrodite individual
Soil organic carbon (SOC) The carbon component of or ganic matter in the soil
   DI SACCO et Al .
biodiversity and ecosystem services, compared with intensive
monoculture plantations;
(iv) Restoration and sustainable management of existing agricultural
land, including through agroforestry, to provide a mix of carbon
sequestration, biodiversity and livelihood benefits and reduce
pressure on native forests;
(v) Protected native non-forest ecosystems (e.g. grasslands, savan-
nas, wetlands).
Here, we build on current evidence and our own experiences
to propose 10 golden rules (Figure 2) to support the delivery of the
native forest elements of the FLR approach (i, ii and iii above), to jointly
increase carbon sequestration and deliver benefits for biodiversity,
ecosystem services and sustainable livelihoods. Agroforestry and
intensively managed plantations are not within the scope of this
These golden rules provide guidance designed to help policymak-
ers, advisors and practitioners of reforestation project s avoid many
of the pitfalls of large-scale tree-planting initiatives that are currently
causing concern. They are in line with the International Principles
and Standards for the Prac tice of Ecological Restoration (Gann et al.,
2019). We use the term ‘reforestation’ in a general sense to refer to
the creation of restored or livelihood native forests by either tree
planting or natural regeneration (NR), where forest formerly occurred
naturally but has been lost recently. High-quality reforestation can
be considered a nature-based solution (NbS) to the problems of bio-
diversit y loss and climate change (Seddon et al., 2020) and, as such,
our rules are allied to the IUCN Global Standard for Nature-based
Solutions and associated guidance (IUCN, 2020), which sets out
criteria to assess whether a proposed NbS addresses a societal chal-
lenge and guides users through aspects of its implementation.
2.1  |  Protect existing forest first
Before planning reforestation, always look for ways to protect existing
forests , including old- and second-growth, degraded and planted forests.
The loss of natural forests continues relentlessly, despite global
efforts to arrest it. In the humid tropics, an average of 4.3 million ha
of old-growth forest was destroyed each year between 2014 and
2018 (NYDF Assessment Partners, 2019). The New York Declaration
on Forestry (NYDF; https://fores tdecl arati aimed to reduce
deforestation by 50% by 2020, while the United Nations Sustainable
Development Goals aimed to end it by 2020. Not only have both
these targets been missed, but tropical deforestation has actually
accelerated by 44% compared with the 13-year period immedi-
ately before the NYDF in 2014 (NYDF Assessment Partners, 2019).
Deforestation on this scale results in huge CO2 emissions (Seymour
& Busch, 2016).
These losses of natural forest are not readily compensated for
by reforestation (Brancalion & Chazdon, 2017; Meli et al., 2017;
Wh e e ler et al . , 201 6) , and nei t h er for e s t p rot ectio n no r rest o r ati o n
should be invoked as a reason to destroy natural areas elsewhere
(Gann et al., 2019). Intact, old-growth forest is a major long-term
carb on sink due to its co mp lex s tr uct ure, la rg e tre es , accum ul at in g
soils and relative resilience to fire and drought (Luyssaert et al.,
2008; Maxwell et al., 2019). The IPCC acknowledges that most
[destroyed] forest ecosystems will take longer than 100 years to re-
turn to the level of biomass, soil and litter pools [found in forest in
an] undisturbed state’ (Aalde et al., 2006). Recovery of ecosystem
services and biodiversity may take centuries, especially the return
of rare or endemic species, which are particularly vulnerable to
dis tu rbance (G ib son et al. , 2011; Rey Be nayas et al., 20 09). Ext inct
species, of course, will never return. Such a steep decline in in-
tact forest also threatens indigenous cultures and human health
(Watson et al., 2018). Large areas of remnant forest, with healthy,
genetically diverse populations of common plant species are es-
sential to supporting reforestation efforts. They provide the seed
rain for NR (Rule 4); a source of seeds, wildings and cuttings for
the production of resilient planting stock (Rule 7); and they pro-
vide habitat for supporting biodiversity, including seed dispersers
and pollinators.
It is therefore vital to protect remaining natural forests—‘pro-
forestation’, sensu Moomaw et al. (2019). Intact, old-growth for-
est is of the greatest value for carbon storage (Maxwell et al.,
20 19 ) an d wi ldli fe (D eer e et al., 20 20) an d shou ld be prio rit ize d fo r
protection. However, second-growth, degraded or logged-over
forest often dominates the remaining forested land (especially
in Southeast Asia; Br yan et al., 2013) and also needs protection
to prevent continued disturbance and further long-term carbon
FIGURE 1 Example of a problematic tree-planting initiative.
In the highly degraded but previously mega-diverse lowlands of
eastern Madagascar, large scale reforestation was carried out in
the 1980s, covering thousands of hectares with the Australian
Grevillea banksia and other non-native species. The initial intention
was to provide communities with a source of firewood. This goal
met with some success, but there were unintended consequences,
such as displacement of croplands and exclusion of native
biodiversity by the introduced species, with such species showing
potential to become significantly invasive (Kull et al., 2019;
Credit: AA)
DI SACCO et Al .
emissions (Maxwell et al., 2019; Reid et al., 2019). If allowed or
encouraged to regenerate (see Rule 5), it will often rapidly re-
cover biomass, resulting in high rates of carbon sequestration,
especially in areas of high water availability (Poorter et al., 2016).
Action at both national and local levels is needed to protect
forests. Persuading governments and corporations to create and
enforce protected areas and legislate against forest conversion can
be effective. For example, Brazil's Soy Moratorium (2006) and Cat tle
Agreement (20 09) achieved some success in reducing soy and cat-
tle-driven deforestation in the Amazon (Nepstad et al., 2014), al-
though they may have displaced forest conversion to the Cerrado
biome, which saw a spike in deforestation in 2011 (Soares-Filho
FIGURE 2 Ten golden rules for a successful reforestation project. The order of the rules matches the order in which tasks should be
considered during project planning and implementation, although some are interdependent and should be considered in parallel. See tex t for
   DI SACCO et Al .
et al., 2014). The first step towards successful protection at the local
level is often identif ication of the drivers of deforestation, among all
stakeholders (Rule 2). Encroachment may be tackled by developing
alternative livelihoods (Rule 10). When fire is a risk, collaborative
communit y groups can take ac tion to raise awareness, organize fire
patrols and install fire breaks, while overgrazing can be reduced by
controlling livestock density, fencing or by instigating cut-and-carry
feeding systems.
2.2  |  Work together
Involve all stakeholders and make local people integral to the project.
The scale and goals of reforestation projects determine their
impact s and therefore affec t who should be involved. For example,
reforestation on smallholder farms can be done without wider stake-
holder engagement being necessary. For large-scale reforestation
projects, engagement of multiple stakeholders is required, to meet
the diverse goals of enhancing rural livelihoods, biodiversity conser-
vation, carbon sequestration, watershed protection and the provision
of other ecosystem services (Erbaugh et al., 2020). A survey of adap-
tive forest management and FLR projects around the world found that
conflicting goals bet ween local communities and project managers
and lack of community involvement were the most commonly cited
causes of project problems or failure (Höhl et al., 2020). Stakeholders
might be direc tly or indirectly affected by a project 's outcomes and
impact s (Erbaugh & Oldekop, 2018) and may include national and
local governments, forestry departments, NGOs, civil society, the
private sector, landowners, farmers and other land users, as well as
universities, botanic gardens, herbaria and other research institutes.
For successful outcomes in both forest protection and reforesta-
tion, it is vital to include local communities from the planning stage
through to delivery and monitoring (Bloomfield et al., 2019). They
are the key to success and have the most to gain from the project.
If their needs are heard and taken into consideration, and they are
informed about the environmental issues the project is address-
ing, they are more likely to support the project and help to deliver
successful outcomes in the long term. Simultaneously, community
provision of labour for forest protection, land preparation, planting
and maintenance provides an opportunity to diversify local employ-
ment, thus improving livelihoods. The realization of multiple positive
outcomes through community engagement has been documented
in Nepal, through community-based forest management (Oldekop
et al., 2019), in the Dodoma and Shinyanga regions of Tanzania,
through the ngitili system that uses traditional local knowledge
and participatory land use planning with the government and other
stakeholders (Duguma et al., 2015), and in several other initiatives
in Madagascar and the Brazilian Amazon (e.g. Dolch et al., 2015;
Douwes & Buthelezi, 2016; Urzedo et al., 2016).
Five levels of community participation in projects have been recog-
ni zed (G a nn et al ., 20 19), ra n gin g f rom weak or pass i ve at Leve l 1 (sim ply
informing stakeholders) to fully active at Level 5 (full support and op-
tional involvement, self-management, benefit sharing and succession
arrangements). Increasing engagement increases positive outcomes,
including equitable distribution of benefits, knowledge, natural cap-
ital, economic sustainability and community well-being (Oldekop
et al., 2019). Reforestation project activities should consistently aim
to actively engage local communities by interactive participation or
self-mobilization, where their vision and objectives for reforestation
are taken into full consideration. Passive participation can lead to com-
munity hostility and disputes over access rights, which may be mani-
festations of underlying or deep-rooted issues, such as conflicts over
land tenure (Agrawal & Redford, 20 09; Chomba et al., 2016).
It is crucial to note that communities are not homogeneous units
(Agrawal & Gibson, 1999). They comprise groups of people differ-
entiated by wealth, ethnicity, gender and other socio-economic
stratifications that have dif ferent power relations and interests in
the reforestation process. For instance, in some countries, men and
women have dif ferent rights to land and trees, which affects those
with insecure rights, mostly women, from effectively participating
in reforestation activities. It is essential to consider those inequal-
ities, as well as conflict s between private, communal and political
interests. Stakeholders’ needs may change over time, so their re-
quests should be re-assessed throughout projects and the strategies
adapted accordingly (Lazos-Chavero et al., 2016).
Sharing of both the costs (in terms of time, labour and money)
and the benefits of reforestation (Rule 10) among all stakeholders
should be agreed upon before the first tree goes into the ground
(Figure 3).
2.3  |  Aim to maximize biodiversity recovery to
meet multiple goals
Restoring biodiversity facilitates other objectives—carbon sequestration,
ecosystem services and socio-economic benefits.
Rather than being an end goal in itself, reforestation is a means
to achieving various goals, typically climate-change mitigation,
FIGURE 3 Ensuring appropriate engagement. In a community-
led reforestation projec t using local indigenous species in eastern
Madagascar, members of the loc al community worked together to
restore areas degraded by fire and over-exploitation (Credit: AA)
DI SACCO et Al .
biodiversity conservation, socio-economic benefits (including food
securit y), soil and hydrological stability and other ecosystem services.
These objectives should be defined beforehand, to allow appropri-
ate project planning, implementation and monitoring (Chazdon &
Brancalion, 2019). Achieving high levels of biodiversit y and biomass,
through the native forest approach, enables multiple outcomes to be
delivered simultaneously. High species and functional trait diversity
enhance productivity, ecosystem resilience and the provision of for-
est products and ecological services to local communities. Restoring
the biodiversity levels and exact species composition of the original
forest may not always be possible, at least initially, due to factors
such as alteration of substrates (e.g. after agriculture and mining),
species extinction, lack of propagation techniques or climate shifts
away from the tolerances of the original species. In such cases, other
native species may be considered to reinstate forest cover, and such
decisions should be made with c aution and be based on sound sci-
ence, to avoid losing locally important species. The ideal achievable
outcome is maximization of natural biodiversity, particularly func-
tional diversity, within current and future climatic and edaphic lim-
itations, while acknowledging that tree species composition may
differ from that of pre-deforestation tree communities.
Forest and landscape restoration allows different objectives
to be prioritized in different landscape zones. However, achieving
multiple objectives means accepting trade-offs (Holl & Brancalion,
2020), and these should be agreed by stakeholders at the start of
projects. It is crucial that the reasons for trade-offs are substan-
tiated, based on sound science and best practices, to achieve the
‘highest and best outcomes’ sensu Gann et al. (2019). While trying to
maximize all the benefit s of projec ts, one essential principle should
be kept in mind: do no harm to local communities, native ecosystems
and vulnerable species.
Wher e th e main goal is timb er produc tion an d/or ca rbon se qu es-
tration, plantations of fast-growing monocultures are widely used.
However, it has been demonstrated that, in the long term, restored
native forests maximize biomass and capture far more carbon while
conserving biodiversit y (Díaz et al., 2009; Lewis et al., 2019).
Socio-economic goals often include the improvement of eco-
nomic conditions for local people, including the poorest communi-
ties. Many projects rely on agroforestry and exotic timber plantations
to meet this objective, but natural, restored and livelihood native
forests deliver economic returns, as well as environmental co-ben-
efits, and should be included in a landscape-wide approach. During
timber production, short harvesting cycles quickly release much of
the stored carbon back into the atmosphere, negating the initial car-
bon sequestration. Low-intensity management of livelihood native
forests, for example through selective extraction, preserves bio-
mass by allowing long-term carbon sequestration and natural veg-
etation succession while also benefitting biodiversity (Crane, 2020;
Hu et al., 2020; Noormets et al., 2015). Alternative livelihood mea-
sures should be supported in the interim period before har vesting,
to avoid the continued conversion of forest with high carbon stocks
elsewhere leading to a net emission of CO2. Biodiverse restored na-
tive forests can provide income through carbon credits, payments
for ecosystem services (PES) and non-timber forest products
(NTFPs; Rule 10).
If the main priority of the project is to conserve biodiversity, it
is important to prioritize areas and select species that maximize this
goal (Rules 4 and 5). Different reforestation approaches, planned at
different levels, can be used: (i) Tree level: plant tree species that are
prioritized for conservation, such as threatened species, or those that
provide resources to target animals (Brancalion et al., 2018) or fungi;
(ii) Ecosystem level: plant or assist the regeneration of species that will
recover the typical composition, structure and functioning of refer-
ence, undisturbed ecosystems (Gann et al., 2019), to maximize habitat
provisi on to a div er sit y of na ti ve spe ci es ; (iii) La nd sc ape leve l: max im iz e
landscape connectivity by creating forested corridors and stepping
stones to link remnant forest patches (Newmark et al., 2017).
Restored native forests can deliver multiple products, such as
food, fibre and medicine, ecosystem services, including watershed
protection, shade and erosion control, as well as recreational, ed-
ucational, spiritual or other cultural benefits. Despite the fact that
these benefits are often recognized, needed or demanded by local
people (Brancalion et al., 2014), they are frequently neglected. The
guidelines in this paper aim to maximize ecosystem services, adding
increased value to any tree-planting or restoration project (Burton
et al., 2018).
2.4  |  Select appropriate areas for reforestation
Avoid previously non-forested lands, connect or expand existing for-
est, and be aware of displacing activities that will cause deforestation
Although reforestation interventions are always implemented at
the local scale, site selection usually involves a multiscale approach.
With the emerging engagement of multilateral and international or-
ga ni zatio ns in tre e-p la nti ng init ia tives (Ho ll & Br anc ali on , 202 0), spa-
tial prioritization decisions can be made at a global scale, but most
restoration initiatives involve an evaluation at the landscape level or
below. Decisions based on considering a combination of historical,
ecological and socio-economic factors at different spatial scales are
the most effective.
Key questions when selecting an area for reforestation are as
(i) Was the area previously forested and is it now degraded? Re-
establishing a species-rich forest in such a place is beneficial for
both biodiversity conservation and carbon sequestration, and
helps fight desertification where this is determined by socio-eco-
no m ic fac tor s (Liu et al., 20 2 0). Refo res t atio n in suc h are as is ge n-
erally highly recommended, and the level of tree cover increase
should be calibrated with the reference values of tree cover
of the target ecosystems, to avoid unintended consequences
for biodiversity and ecosystem services. However, in some
previously forested areas, for example South Central United
States oak forest, climate change may drive a transformation
   DI SACCO et Al .
to non-forest biomes, such as savanna and grassland (Millar &
Stephenson, 2015). Modelling tools are needed to evaluate po-
tential target areas and identify those that are approaching such
(ii) Has the area been occupied historically by a non-forested biome
such as grassland, savanna, non-forested wetland or peatland?
Afforestation in such areas depletes both biodiversity and SOC
(Bond et al., 2019; Friggens et al., 2020; Veldman et al., 2015)
and must be avoided. For example, grasslands often host high
biodiversity and many threatened species, as well as contributing
significantly to belowground carbon sequestration (Burrascano
et al., 2016; Dass et al., 2018). Non-forested peatlands contain
an even higher amount of SOC, which would be released into the
atmosphere if trees were planted there (Brancalion & Chazdon,
2017; Crane, 2020; NCC, 2020). Similarly, lands covered by snow
at high latitudes reflect an important quantity of sun radiation
due to the high albedo, providing a cooling effec t on the planet
that would not be compensated for by the amount of carbon
slowly captured by trees grown in those cold climates (Bala et al.,
2007; Betts, 2000). A critical step for tree-planting initiatives is
therefore to define ‘no-go zones’, where restoration should focus
instead on non-forest vegetation;
(iii) What are the wider effects of refores tation in the target area, includ-
ing impacts on groundwater, biodiversity, climate, ecosystem ser-
vices and livelihoods? If the area is dry and water is scarce, trees
could reduce the groundwater and river flow, with negative con-
sequences for local inhabitants (Allen & Chapman, 2001; Feng et
al., 2016). However, in seasonally dr y climates, restoring forests
on degraded watersheds can help to increase water infiltration
and reduce surface run-off during the rainy season, reducing ex-
treme fluctuations in streamflow throughout the year (Gardon
et al., 2020). In urban areas, trees can be planted to mitigate the
direct effects of GCC, providing an additional contribution to the
carbon sequestration needed (Parsa et al., 2019) while also de-
livering other ecosystem services such as the provision of recre-
ational spaces, wildlife habitats, clean air and shade;
(iv) How close is the land to areas of natural forest? This affects both
the capacity of the site to regenerate naturally (Rule 5) and the
value of the reforested site to biodiversit y, for example by creat-
ing buffer zones, corridors and stepping stones enabling native
species to migrate bet ween forest remnants and expand their
distribution (Tucker & Simmons, 2009);
(v) Who is currently using the land, how will they be compensated
for any income losses and where will they move their activities? If
these factors are not considered, the land might be retaken sub-
sequently, or further deforestation or social conflict s might be
caused elsewhere (Cuenca et al., 2018; Mey froidt et al., 2010).
Issues of land tenure and forest governance are critical to the
success of reforestation and are safeguarded in the Cancun
Agreement (UNFCCC, 2011). Protecting and restoring degraded
forest remnants is the best way to increase carbon stocks and
decrease habitat fragmentation without using non-forested land
that may already be in use (Brancalion & Chazdon, 2017).
More tools and tailored resources are needed to help guide these
decisions. The Restoration Opportunities Assessment Methodology
(IUCN, & WRI, 2014), for instance, has been used in many countries
that have made pledges to the Bonn Challenge, to identify FLR op-
portunities. The resulting maps identify high-priority areas for in-
tervention and provide a helpful framework for determining what
method is best. Technologic al advances will provide new tools and
resources, such as NASA's Global Ecosystem Dynamics Investigation,
which will facilitate the use of LiDAR to prioritize areas of degraded
forest for restoration (Deere et al., 2020).
2.5  |  Use natural regeneration wherever possible
Natural regeneration can be cheaper and more effective than tree plant-
ing where site and landscape conditions are suitable.
The NR approach to forest restoration spans a spectrum of dif-
ferent levels of human intervention:
(i) No inter vention or passive restoration (Chazdon & Uriate, 2016);
(ii) Low inter vention, including protection from further damage
such as grazing or fire, and rewilding, which includes the selec-
tive reintroduction of missing fauna to restore natural processes
(Perino et al., 2019);
(iii) Intermediate intervention, including enrichment of naturally
regenerated forest by selective planting of missing species and
assisted NR (ANR; FAO, 2019), where weeds are cleared around
naturally regenerating trees to accelerate their growth. ANR
has been used to restore Imperata grasslands in the Philippines
(Shono et al., 2007) and logged-over forest that has become
dominated by lianas (Philipson et al., 2020);
(iv) High intervention, including the framework species approach
(Rule 6) and applied nucleation (Za hawi et al., 2013), wh er e pa rts
of the site are intensively planted to facilitate NR in the rest of
the site.
When carbon capture and biodiversity enhancement are primary
objectives, NR can provide significant benefit s over tree planting, if
practised in suitable locations, as described below. Carbon seques-
tration in naturally regenerated areas is potentially 40 times greater
than in plantations (Lewis et al., 2019) and species richness is gener-
ally higher, particularly for forest specialist species (Barlow & Peres,
20 0 8; Br ocker h of f et al., 20 08; Roz end aal et al. , 20 19). NR is also sig-
nificantly cheaper than tree planting, with studies in Brazil showing
implementation costs reduced by 38% (Molin et al., 2018) or even up
to 76% (Crouzeilles et al., 2019). However, this approach is unsuit-
able for certain ecosystems, for example those in ‘old, climatically
buffered infertile landscapes’ (‘OCBIL s’, sensu Hopper, 2009) found
in biodiverse regions, such as the southwestern Australian biodiver-
sity hotspot. In such landscapes, natural recolonization processes
are incapable of reinstating ecosystems once the native vegetation
has bee n removed , an d substa ntial rep la nting and see di ng are there-
fore required (Koch & Hobbs, 2007).
DI SACCO et Al .
Once a land area has been targeted for natural or seminat-
ural forest cover, the two key questions are as follows: (i) Is
the forest capable of returning spontaneously? and (ii) What level
of intervention is required to assist and accelerate the regenera-
tion? Th e site's potential fo r N R will depend on mult iple fac tors ,
which can be considered at the landscape and site level (Elliott
et al., 2013).
At the landscape level, the first step should be to identify and
control the factors that led to deforestation in the first place—a
task that should involve all stakeholders (Rule 1). One of the most
important landscape fac tors is the proximity of the site to areas of
remaining natural forest that can serve as a diverse source of nat-
urally dispersed seeds. Crouzeilles et al. (2020) found that 90% of
passive regeneration occurred within 192 m of forested areas, while
Molin et al. (2018) found best results within 100 m of the near-
est forest. The presence of birds and animals in and around a site
is crucial for seed dispersal of many plant species. Typically, large
wild animals and birds are the first to be locally extirpated, in which
case the plants they dispersed may fail to recolonize unless manu-
ally introduced (enrichment planting). Another key factor is climate,
particularly mean annual precipitation (Becknell et al., 2012). In the
Neotropics, biomass recovery in second-growth forests was up to
11 times higher in wetter areas (Poorter et al., 2016).
At the site level, the previous land use and degree of degrada-
tion affect the regeneration potential, with heavily degraded sites
(e.g. former mine sites) invariably requiring active interventions
such as planting and topsoil replacement (Meli et al., 2017). The
size of the target area will clearly affect distance to the nearest
fo re s t (an d t hu s the re gen era tio n pot ent ial of lig htl y or mod e ratel y
degraded sites), with central parts of the site being further away
than the nearest edges. Different levels of intervention may there-
fore be required within a single large site.
The exi st ing natur al vege ta tion currently present on a site has
the most immediate effect on determining the regeneration path-
way. In a lightl y deg raded site, a dense commun it y of tre e stu mp s,
seedlings and a diverse soil seed bank enable rapid regeneration,
especially in humid tropical areas, potentially achieving canopy
closure in under a year (Elliott et al., 2013). Advice on the re-
quired densit y of regenerants for NR ranges widely from 200/ha
(Shono et al., 2007) to 3100/ha (Elliott et al., 2013) and depends
on climate. The stocking density required to achieve rapid canopy
closure is lower in warm wet climates, since tree crown expan-
sion oc cu r s m or e rapi dly th an in co ol, dr y clim at es. He r ba ceo us or
woody weeds usually out-compete regenerating trees and should
be controlled through cutting, pressing or ‘lodging’ (flattening
weeds with a board), mulching, herbicides or controlled grazing,
that is, through ANR (FAO, 2019).
Other important site factors are soil quality, topography and
hydrological features (Molin et al., 2018). Given the complex in-
teraction of all thes e fa ctors, the best way to deter min e th e site's
suitability for NR and the level of human inter vention required
is to take an experimental and adaptive management approach
(Rule 9).
2.6  |  Select species to maximize biodiversity
Plant a mix of species, prioritize natives, favour mutualistic interactions
and exclude invasive species.
Tree planting is needed to restore forest when NR is insufficient
(Rule 5). The International Standards for Ecological Restoration spec-
ify a ‘native reference ecosystem’ to guide species selection (Gann
et al., 2019). In heavily degraded sites, species should be selected
based on their ability to est ablish in altered or unfavourable condi-
tions, which might include compacted soil, drought and competitive
weeds. Native pioneer species are most likely to survive initially,
while late successional species can be intercropped with these pio-
neers, be introduced with successive planting interventions or may
even eventually colonize the site naturally.
The framework species approach to forest restoration in the
tropics is a highly effective tree-planting option that depends on
the selection of a suite of native species with specific functional
traits (Goosem & Tucker, 2013). It involves planting the fewest trees
needed to complement and promote NR and recapture the site from
weeds in 2–3 years. Framework tree species are characteristic of the
reference ecosystem and have: (i) high survival and growth rates; (ii)
dense, spreading crowns that shade out herbaceous weeds; and (iii)
traits that attract seed-dispersing wildlife (e.g. flowering/fruiting at a
young age). Mix tures of 20–30 species (both pioneer and climax tree
species) should be planted. Biodiversity recovery depends on rem-
nants of the reference forest type occurring within a few kilometres
of the restoration site (as a seed source) and seed-dispersing animals
remaining in the landscape (Elliott et al., 2013). A successful case of
fra mework spe cie s appro ach applie d in Thailan d is shown in Fig ure 4.
Maximizing biodivers ity depends not onl y on the numbe r of sp e-
cies reintroduced but also on the functions they perform. Promoting
mutualistic interactions, such as those involving native tree species
and fungi, seed-dispersing animals, pollinators and other organisms,
is crucial to achieving a resilient, biodiverse restored ecosystem
(McAlpine et al., 2016; Steidinger et al., 2019), but the impor tance of
such interactions is often underestimated.
Rare, endemic or threatened taxa are less likely to colonize
through natural succession (Horák et al., 2019) and should there-
fore be reintroduced at the appropriate stage of forest maturity. This
practice will contribute to the survival and conservation of the most
vulnerable species. Such species can contribute greatly to carbon
stocks, since they tend to be late-successional species with dense
wood (Brancalion et al., 2018).
The GlobalTreeSearch database (
tree_search.php) lists all known tree species and can generate
checklists of native species for each country. Local specialists
including botanical experts and restoration ecologists should be
consulted, to determine which native species are most suitable
for the particular forest type being restored. The Global Tree
Assessment (www.globa ltree asses aims to deliver tree
conservation assessments for all tree species by the end of 2020.
This will help identify threatened species that can be included in
restoration projects.
   DI SACCO et Al .
In livelihood native forests, selecting a mix of species, rather than
planting a monoculture, is crucial (Brancalion & Chazdon, 2017). A
mixed-species forest, either with native species only or with a mix of
native and non -n at iv e sp ecies , ha s a hi gh er ca pa ci t y to con se r ve bio -
diversit y, create habitats for wildlife and at tract seed dispersers and
pollinators. Such forest can regenerate autonomously, especially if
patches of native vegetation are maintained within the plantation
matrix as habitat islands (Horák et al., 2019). It will also be more re-
silient to disease, fire and extreme weather events (Florentine et al.,
2016; Verheyen et al., 2016). Monoculture plantations sequester lit-
tle more carbon than the degraded lands on which they are planted,
especially if they are used for fuel or timber, in which case carbon
is released back into the atmosphere within a few decades (Körner,
2017; Lewis et al., 2019).
Including exotic species in livelihood native forests is controver-
sial (Catterall, 2016). For example, eucalypts (Eucalyptus) may have
high cash value, but eucalypt plantations suppor t lower biodiversity
than native forest s (Calviño-Cancela et al., 2012) and are colonized
by mainly generalist plant and animal species (Brockerhoff et al.,
2008). A major concern is that exotic species often become inva-
sive, for example certain Australian Acacia species in South Africa
(Richardson & Kluge, 2008). Invasive species rank second only to
habitat loss and degradation as a cause of the current global biodi-
versity crisis (Bellard et al., 2016). They have long-term effects on
the environment, compete with native species, reduce biodiversity
and often reduce water availability (Dyderski & Jagodziński, 2020;
Scott & Prinsloo, 20 08). Their removal, which needs to be done be-
fore restoration interventions can commence, is invariably difficult
and very expensive. Invasive exotic species should never be planted.
However, under certain circumstances, some exotic, non-invasive
species can be good allies for tropical forest restoration. In a humid
tropical region of Brazil, exotic eucalypt s, when planted in mixed plan-
tations with native species and selectively harvested after 5 years,
allowed the NR of native trees in the understorey and substantially
defrayed restoration costs (Brancalion et al., 2020). Crucially, the eu-
calypt s did not regenerate from seed.
Further research is required to identify more high-value native
species that could be used instead of, or together with, desired ex-
otic species. For example, in Kenya, Melia volkensii is a popular nat ive
timber species and has a lower water demand than exotic eucalypts
(Ong et al., 2006; Stewart & Blomley, 1994). The use of mainly na-
tive species in new livelihood native forests has been successful
in Latin and Central America, where companies such as Symbiosis
Investimentos and Sucupira Agroflorestas are developing propaga-
tion protocols for native species, promoting agroecological princi-
ples, practising sustainable forestry, and in some cases conserving
and restoring natural forest alongside plantations.
Adaptability to GCC should also be considered when selecting
species for both native and livelihood native forests. When GCC is
proven to negatively impact native species, non-native species could
be considered on the basis of preserving ecosystem functions. Such
species must be subjected to comprehensive risk assessments that
include biosecurity threats and potential invasiveness (Ennos et al.,
2019). This could form part of an assisted migration programme.
2.7  |  Use resilient plant material
Obtain seeds or seedlings with appropriate genetic variability and prov-
enance to maximize population resilience.
To ensure the survival and resilience of a planted forest, it is vital
to use material with appropriate levels of genetic diversity, consis-
tent with local or regional genetic variation. Vegetative propagation
or using seeds with low genetic diversity generally lowers the resil-
ience of restored populations through reduced evolutionary poten-
tial and problems with inbreeding depression (Thomas et al., 2014).
As a result, planted forest s may be disease-prone and unable to adapt
to long-term environmental change. Such genetic bottlenecks can
result from poor seed-collection strategies, such as collecting from
too few parent trees or declining source populations. Using material
from well-designed seed orchards, or, in the many cases where this
is not available, mix tures of seed with different provenances, usually
FIGURE 4 Example of successful tree planting. The framework species method of forest restoration can be effective even on the most
degraded sites, provided intact forest remains nearby. (a) August 2012: Siam Cement's limestone mine in Lampang Province, northern
Thailand. (b) April 2013: after spreading the site with topsoil (60 cm deep), 14 framework tree species were planted, including several Ficus
species and native legumes, to improve soil conditions. Corrugated c ardboard mulch mats were also applied. (c) February 2015: by the end
of the third rainy season, canopy closure was achieved and macaque monkeys star ted visiting the plot to eat figs, in the process naturally
dispersing seeds of other species through defecation. Mean sur vival across species was 6 4% and relative annual growth rate averaged 91%
(Credit: Siam Cement Group and SE)
(a) (b) (c)
DI SACCO et Al .
increases genetic diversity in planted forests (Ivetić & Devetaković,
2017). However, in exceptional ecosystems, such as Australian and
African OCBILs, which have strong local adaptation, (Hopper, 2009;
James & Coates, 2000), highly local provenancing may be required.
Best practice involves collecting seeds from many individuals
across the full ex tent of the parent population randomly, to include
the rarest alleles (Hoban, 2019; Hoban & Strand, 2015). Similarly,
Ivet and De ve tak ov ić (2017 ) ide nt ifi ed th e si ze of the par ent al po p-
ulation as a key determinant of genetic diversit y in planted forests;
they viewed provenance and seed-collecting strategies as the most
imp ort ant ma na ge me nt pra ctices in tree-p la nt in g pr oj ec t s. As a ge n-
eral rule, for adequate genetic diversity, seed should be collected
from at least 30 individuals of outcrossing species and at least 50
individuals of selfing species (Pedrini, Gibson-Roy, et al., 2020).
Seed collection from local parent populations is advised since
genotypes are adapted to climatic and environmental conditions
similar to those of restoration sites. However, more dist ant prov-
enances may be considered if conditions are similar across a large
part of a species’ range, or to match conditions under future GCC
scenarios (predictive provenancing). If decisions are being made
based on climate predictions, then sound science and experimen-
tal eviden ce of why cl im ate-a da pt ed gen et ic mat er ia l is being used
should be ar ticulated (Alfaro et al., 2014). A cautious strategy is
to use composite provenancing sensu Broadhurst et al. (2008).
Seed zone maps can help practitioners to identify appropriate
provenances of material for planting target sites; however, such
maps are rare for most forest systems, particularly for understorey
One of the main bottlenecks for forest restoration is inade-
quate supply of native plant material. Lack of see ds (Jalo nen et al.,
2018; León-Lobos et al., 2020; Merritt & Dixon, 2011) and plant-
ing stock (Bannister et al., 2018; Whittet et al., 2016) of target
species from appropriate sources in the required amounts are
often critically limiting. This problem is particularly acute in the
tropics, where many tree species produce seeds that are difficult
to store (i.e. are desiccation sensitive) and for animal-dispersed,
large-seeded tree species, which are of crucial importance for
forest restoration (Brancalion et al., 2018). In addition, many of
the seed supply sources are forestry genebanks that often have
different aims, such as conserving desired traits rather than broad
genetic diversity.
2.8  |  Plan ahead for infrastructure, capacity and
seed supply
From seed collection to tree planting, develop the required infrastruc-
ture, capacity and seed supply system well in advance, if not available
externally. Always follow seed quality standards.
For project s involving tree planting or direct seeding, appropri-
ate infrastructure and seed supply systems are essential. Decisions
should be made at least a year in advance on whether to source
seeds and produce seedlings in-house, subcontract these tasks or
purchase plant material from external suppliers. If seeds are pur-
chased externally, suppliers should be able to provide information
on seed quality and the legality of their collec tion (Pedrini & Dixon,
2020). If commercial suppliers of seeds and seedlings fail to meet
project requirements for species mix, quantity, genetic diversity,
provenance or quality (Rule 7), projects may need to develop their
own collection, storage and propagation capacity.
Where seed is self-sourced, national legislation and local laws
on access to biological material (UN Convention on Biological
Diversit y, 2011) ( and international seed stan-
dards (e.g. ENSCONET, 2009b; Pedrini & Dixon, 2020) must be
followed, to ensure seeds are high quality and to avoid damag-
ing source populations by over-collecting (no more than 20% of
the available ripe seeds should be collected). Basic equipment for
wild-seed collecting, cleaning and storage is needed. Collecting
from tall trees requires specialist equipment, including extendible
pruners, throw lines, tarpaulins and tree-climbing harnesses. Seed
collectors should be trained to use this equipment efficiently and
safely. Training should include phenological monitoring and seed
physiolog y, to ensure that collecting trips are timed efficiently at
peak fruiting times and when maturity is optimum (Kallow, 2014).
Involving botanist s and local experts enables species identifica-
tion, efficient location of trees of target species and optimum
timing for collection. Data on species identification, ecological
conditions and provenance should be recorded simultaneously
with the seeds. Alternatively, seeds can be provided by a third
party, either collected directly from the wild or from wild-origin
seed orchards, usually by state agencies or commercial suppliers
(Pedrini, Gibson-Roy, et al., 2020).
If collecting seeds, the seed storage behaviour of the target
species should be checked first, so they are handled appropriately.
Orthodox seeds can be stored in seed banks, increasing their lon-
gevity for decades and allowing their use over extended periods,
which optimizes collecting efforts and reduces waste (De Vitis et al.,
2020; ENSCONE T, 2009a). Literature on seed storage behaviour is
available for many taxa (Hong et al., 1998), and it is possible to pre-
dict (Wyse & Dickie, 2018) or test (Hong & Ellis, 1996; Mattana et al.,
2020) the behaviour of understudied species. The Seed Information
Database,, curated by the Royal Botanic
Gardens, Kew (RBG Kew), stores information on a wide range of
Low-cost seed-storage facilities can be installed if seed banks
are not available regionally. Further information from RBG Kew is
freely available here: http://brahm sonli ing/
Resou rces. Seed banking is particularly useful in arid and semi-arid
biomes (León-Lobos et al., 2012), where over 97% of the species are
estimated to have orthodox seeds, but it is also a valuable option for
the majority of species in humid ecosystems (Wyse & Dickie, 2017).
Propagation protocols are available for many common spe-
cies, but if they are not, then germination trials are required. The
seeds of most wild species have dormancy mechanisms (Baskin
& Baskin, 2014), requiring specific conditions for germination.
These can sometimes be deduced from the seed morphology and
   DI SACCO et Al .
ecology of each species (Kildisheva et al., 2020), but empirical re-
search may be required to achieve germination for species with
deep dormancy.
If direct seeding is chosen, then seed priming (for optimal germi-
nation) and/or coating (to protect seeds from predators, desiccation
and diseases) is beneficial (Madsen et al., 2012; Pedrini et al., 2020;
Williams et al., 2016). The number of seeds required is much higher
than the target number of trees, since conversion rates of seeds to
establi shed seedli ngs are usu ally ve ry low and are high ly specie s-de-
pendent (James et al., 2011) and site-dependent (Freitas et al., 2019).
The development of a seeding plan that includes site preparation
and seeding strategy, as well as monitoring after planting, is crucial
for success (Shaw et al., 2020), while adopting emerging technolo-
gies can help to optimize seed use efficiencies (Pedrini, Dixon, et al.,
If saplings are to be planted, an in-house nursery must be built
(Elliott et al., 2013) or an appropriately accredited nurser y selected
for their production. If such infrastructure and expertise are not
available locally at the start of the project, it is import ant to include
them in project planning. Local people are import ant as sources of
both labour and expertise. Opportunities to convert private agricul-
tura l or horticu lt ural fa cilitie s in to the re so urces nee ded for the proj-
ect should be explored.
2.9|  Learn by doing
Base restoration interventions on the best ecological evidence and indig-
enous knowledge. Perform trials prior to applying techniques on a large
scale. Monitor appropriate success indicators and use results for adap-
tive management.
Planning decisions should be made by combining both scientific
and indigenous knowledge. Traditional knowledge, acquired over
many generations by people who have lived close to the forest, is
particularly useful where field experiments to generate scientific ev-
idence may take a long time to yield results (Wangpakapat tanawong
et al., 2010). International standards (e.g. Gann et al., 2019) give
general guidance, while Floras, previous project report s and the sci-
ent ific literature can pr ovide more specific infor mation su ch as func-
tional trait data to aid species selection (Chazdon, 2014).
Ideally, small-scale trials should be implemented before large-
scale tree planting commences, to guide species choices and test the
effectiveness of proposed techniques. These may include land man-
agement interventions to overcome site-specific barriers, such as
degraded soils (Arroyo-Rodríguez et al., 2017; Estrada-Villegas et al.,
2019), competitive weeds (FAO, 2019), fire and herbivores (Gunaratne
et al., 2014; Rezende & Vieira, 2019), and the absence of mutual-
istic organisms in soils, such as mycorrhizal fungi (Asmelash et al.,
2016; Fofana et al., 2020; Neuenkamp et al., 2019). Unfortunately,
trials take years to yield results, so projec ts often have to be initiated
through the exchange of previous knowledge. Subsequent monitor-
ing then generates data for adaptive management, a fundamental
principle of FLR since its inception (Gilmour, 2007).
For monitoring forest restoration sites, it is useful to establish
permanent sample plots in: (i) the restoration site (treatment); (ii)
a site where no interventions are implemented (control); and (iii) a
reference forest remnant (target). Comparing (i) and (ii) determines
the effectiveness of restoration interventions. Comparing (i) and
(iii) tracks the progress of restoration towards the target end-
state. Data should be collected before and just after restoration
interventions are initiated (baseline) and annually thereafter, at
least until canopy closure.
Restoration progress is indicated by the biomass, forest struc-
ture, biodiversit y and ecosystem functioning in restoration sites
all trending towards those of the reference (or target) ecosystem.
However, monitoring can focus on biomass and biodiversity, since
the other two ecological indicators and many socio-economic bene-
fits (Table 2) stem from them.
TAB LE 2  Why income-generating forest ecosystem services increase with both biomass accumulation and biodiversity recovery (both of
which are higher in existing and restored native forests than in monoculture plantations)
ecosystem ser vice Biomass accumulation Biodiversity
Carbon storage About half (~47%) of all tree biomass is carbon1Biodiversity increases biomass accumulation2
Forest products Biomass accumulation increases the quantity of
Biodiversity increases the variety of produc ts, providing
economic security against fluctuating market prices
Watershed services I:
Flow regulation (flood/
drought mitigation;
irrigation for
Biomass accumulation increases organic matter
accumulation and thereby soil moisture-holding
Tree species diversit y incre ases interception, decreases
runoff (flash floods) and improves inf iltration3
Watershed services II:
Soils (erosion landslide
Biomass accumulation increases belowground root
biomass and thereby reduces erosion and landslides
Different tree species root to dif ferent depths, decreasing
Ecotourism Biomass accumulation increases ecosystem structure,
niches and biodiversity
Biodiversity-rich native forests att ract more ecotourist s
Sources: 1Mar tin and T homas (2011); 2Steur et al. (2020); 3Gardon et al. (2020).
DI SACCO et Al .
Biomass is estimated from stocking density and tree sizes in
sample plots. Allometric equations are used to derive biomass and
carbon from measurement s of tree diameter and height, and wood
density (Chave et al., 2014). Soil samples should also be collected to
determine soil carbon. Ground surveys are rapidly being replaced
by aerial photogrammetry (de Almeida et al., 2020) using drones to
create 3D forest models, within which the heights and shapes of all
trees can be measured. However, to gather species-specific data
and calibrate remote sensing approaches, ground surveys remain
It is impractical to monitor all species to assess biodiversity re-
covery, so biodiversit y indicator groups are used, most commonly
plants and birds. For trees and ground flora, the abundance of spe-
cies in sample plots should be recorded and the data used to con-
struct species-effort curves and calculate diversity indices (Ludwig
& Reynolds, 1988). To monitor bird species richness, we recommend
the Mackinnon List Technique (Herzog et al., 2002). If resources are
available, more comprehensive biodiversity assessments using envi-
ronmental DNA and insect traps can provide rich and cost-effective
data (e.g. Ritter et al., 2019).
Monitoring should also assess progress towards project-specific
goals, such as erosion control or recovery of an endangered species.
Where livelihood benefits are a key objective, they may be assessed
using indicators such as jobs created or changes in income, and equity
in distribution at the gender, household and communal levels. Where
income is to be generated from extraction of timber or NTFPs, it is
vital to achieve sus tainable prod uction by ensuri ng th at harvest rates
of products do not exceed their productivity. This can be monitored
through simple ‘yield-per-unit-effort’ techniques—recording product
quantities harvested and harvesting time expended—with communi-
ty-agreed reductions in har vesting intensity, if yields start to decline.
Monitoring and verification of restoration, particularly to claim
income from carbon credit s and other environmental services, is
usually carried out by independent assessors at great expense.
However, studies have shown that local people are capable of per-
forming monitoring more cost-effectively (Boissière et al., 2017;
Danielsen et al., 2013), and their indigenous knowledge is of great
value to the process ( Wangpakapattanawong et al., 2010).
2.10|  Make it pay
Develop diverse, sustainable income streams for a range of stakeholders,
including carbon credits, NTFPs, ecotourism and marketable watershed
Income generation by selling forest products from livelihood
forests is easily achieved, whereas marketing environmental ser-
vices from existing and restored native forest is more difficult,
particularly in protected areas. However, the sustainability of
forest restoration depends on income streams generated from it
exceeding those from alternative land uses and on that income
being shared fairly among all stakeholders, including the poorest
(Brancalion et al., 2012).
In 2009, The Economics of Ecosystems and Biodiversity initia-
tive estimated the value of tropical forest ecosystem ser vices to be
USD 6120/ha/year (USD 7732 today, after inflation), based on data
from 109 studies (TEEB, 2009). Watershed ser vices contributed
most (38.8%), followed by climate regulation (32.1%), provisioning
services (21.5%) and recreation/tourism, (6.2%). All these values
depend on the two fundamental indicators of restoration: biomass
accumulation and biodiversity recovery (Table 2).
REDD+ has made some progress with monetizing forests as car-
bon sinks (Angelson et al., 2012). Forest carbon value alone often
exceeds revenue from the main drivers of deforestation (e.g. oil
palm; Abram et al., 2016), but application of REDD+ to incentiv-
ize restoration has been problematic, due to issues of governance
and socio-economic conditions, particularly fluctuations in carbon
credit prices. To ensure revenue flows mostly into local economies,
local people should have direct access to carbon markets as well as
low-interest start-up loans, to fund restoration work and support
their families until break-even is achieved. Fur thermore, transac tion
costs, including monitoring, reporting and verification, should be
minimized by building local capacity, to reduce dependency on paid
external agents (Köhl et al., 2020).
While NTFPs are usually less valuable than carbon, local peo-
ple can easily monetize them, and start-up investment is minimal (de
Souza et al., 2016). Furthermore, NTFPs can provide security and
adaptability during periods of financial hardship (Pfund & Robinson,
2005), and their diversity buffers against fluctuating markets—if the
price of one product falls, another can be substituted. Conversely,
monoculture plantations leave farmers vulnerable to fluctuations in
a single commodity market price. Thus, biodiversity recover y drives
both ecological stability and economic security. However, to ensure
sustainable production, harvesting rates must be sustainable and
therefore monitored (Rule 9).
Watershed ser vices are the most difficult to monetize, since they
constitute ‘avoided detrimental impacts’, such as flood damage or
decline of agricultural productivity. The need for such services is un-
predictable in time and place. They are a ‘public good’, rather than a
readily quantifiable commodity. Consequently, government funding
(via taxes or water charges) is the most appropriate monetization
mechanism. Several such schemes have been well documented in
Latin America and China (Porras et al., 2008).
Ecotourism can be a lucrative source of loc al income, which di-
rectly monetizes biodiversity. However, its potential is often over-
estimated. Substantial start-up funding is needed, particularly for
accommodation construction. Furthermore, the skilled labour re-
quired to meet the discerning demands of ecotourists is often im-
ported from outside, sidelining local people.
Innovative marketing will be essential, to turn restoration values
into financial incentives, since both investors and the public are unfa-
mi l iar wi t h pay i ng fo r some of th e ser vice s outli n ed abo ve (B ran c a lio n
et al., 2017). Comprehensive socio-economic monitoring will also be
needed, to ensure that payments actually benefit local communities
and that changes in land and resource values have no deleterious
social consequences. Finally, if such financial incentives lead to a
   DI SACCO et Al .
surge in restoration project s at the expense of agriculture, the prices
of carbon credits and NTFPs could crash and food production could
decline, resulting in increased food prices and reduced food security.
Models of the potential macro-economic effects of restoration fi-
nancing are therefore also needed, to forestall such impacts.
The guidelines presented here show that reforestation is more com-
plex than is of ten initially thought. There is no universal, easy solution
to a successful initiative given the extraordinar y diversit y of species,
forest types, sites, and cultural and economic environments. In many
cases where livelihoods depend upon altered landscapes, restoration
goals can only be achieved through creating a mosaic of land uses at
the landscape level and by engaging with societ y at large (Figure 5).
Despite the inherent complexity of reforestation initiatives, there
are succes sful exa mpl es to bu ild on and develo p fur ther. Over the past
30 years, ecologists have transformed the concept of forest restoration
to an attainable goal, having developed tools to overcome the technical
and knowledge barriers to its implementation through robust scientific
research. This means that calls by the UN and many other organiza-
tions to restore forest to hundreds of millions of hectares worldwide—
inconceivable before—are becoming increasingly feasible. However,
achieving such ambitious goals will only happen through careful con-
sideration of the diverse aspect s discussed in this review.
Partnerships involving multiple stakeholders (corporates, gov-
ernments, NGOs, scientists, practitioners, landowners) are likely
to yield the most enduring long-term benefits. Overcoming the so-
cio-econ omic and poli ti ca l barriers to for es t re storati on will als o re -
quire good governance, long-term funding mechanisms, enshrined
legal protective measures for the restored sites, and effe ctive com-
munication among stakeholders at the science–policy–practice
Vast reforestation programmes are now underway across the
planet, and these will require monitoring so that learning opportu-
nities are not lost. We need to rely on the best scientific evidence
available and implement carefully planned, replicated, controlled
FIGURE 5 Schematic view of a successful reforestation programme. This landscape contains several components: (a) protected existing
native forests, either old- or second-growth, where native seeds are collected; (b) restored riparian forest creating a biological corridor
connecting remaining forest patches; (c) a naturally regenerating area, adjacent to an existing native forest that provides seed rain for
natural regeneration; (d) restored or livelihood native forest, which might include non-invasive exotic useful species for timber and non-
timber forest products (NTFPs), where people monitor biomass and biodiversity recovery; (e) tree nursery and seed bank where native
seeds are stored and propagated; (f) tree planting area, with a section dedicated to establishment trials; (g) protected native non-forest
ecosystems, such as grassland and wetland; (h) urban and rural areas, with sustainable agriculture and livestock
DI SACCO et Al .
experiments on large spatial scales. This is key to objectively testing
and continuously improving the effectiveness of existing socio-eco-
nomic constructs, such as community forestr y, REDD+, FLR and
PES. Crucially, politicians and policymakers need to act now to en-
gineer a rapid paradigm shift in the way we protect existing forests
and restore new ones using native species, to benefit both people
and nature. They should use innovative regulations, incentives and
all the levers at their disposal.
The massive reforestation initiatives currently underway, the up-
coming UN Decade on Ecological Restoration and aspirations for a
post-COVID green recovery, have generated unparalleled hope and
optimism that forest restoration really can improve global ecology
while uplifting local livelihoods. However, it will only do so if it is
based on sound science, guided by indigenous knowledge and local
comm un it ies , su ppo rte d by fai r go ve rn ance, and incentiv iz ed by lo ng-
ter m funding mec hanisms . We hope that the 10 golden rul es outlined
here will help guide all those who are involved in restoring Earth's
forests to address such issues fruitfully and to turn the hope and op-
timism into reality.
We thank the editor and t wo anonymous reviewers for construc-
tive feedback that helped improve this manuscript. We also thank
Jill Kowal, Laura Kelly, James Borrell, Tiziana Ulian and several
other colleagues for input and discussions. We acknowledge fund-
ing from Sk y Zero to A .D.S.; Chiang Mai University, Thailand to S.E.;
the Swedish Research Council (grant # 2019-05191), the Swedish
Foundation for Strategic Research (grant # FFL15-0196) to A.A.; the
Royal Botanic Gardens, Kew to A. A., E.B., A.D.S., K. A.H. and R. J.S.;
the São Paulo Research Foundation (FAPESP, grant #2018/18416-2)
to P.H.S.B and the European Union to the Regreening Africa pro-
gramme which supported S.C. The funders had no influence on the
research reported here. The authors declare no competing interests.
Alexandre Antonelli, Kate A. Hardwick, Paul Smith and Alice Di
Sacco conceived the initial outline of the article; Alice Di Sacco and
Kate A . Hardwick led the writing, coordinated the author contribu-
tions and prepared the figures; all authors contributed to writing,
developing and reviewing the manuscript.
Data sharing not applicable—no new data generated.
Alice Di Sacco
Kate A. Hardwick
Pedro H. S. Brancalion
Elinor Breman
Loic Cecilio Rebola
Susan Chomba
Kingsley Dixon
Stephen Elliott
Paul Smith
Rhian J. Smith
Alexandre Antonelli
Aalde, H., Gonzalez, P., Gytarsky, M., Krug, T., Kurz, W. A., Ogle, S.,
Raison, J., Schoene, D., Ravindranath, N. H., & Nagmeldin G. E.
(2006). Chapter 4. Forest lands. In H. S . Eggleston, L. Buendia, K.
Miwa, T. Ngara, & K. Tanabe (Eds.), 2006 IPCC guidelines for national
greenhouse gas inventories. Volume 4: Agriculture, forestry and other
land use. IGES.
Abram, N. K., MacMillan, D. C., Xofis, P., Ancrenaz, M., Tzanopoulos, J.,
Ong, R., Goossens, B., Koh, L. P., Del Valle, C., Peter, L., Morel, A. C.,
Lackman, I., Chung, R., Kler, H., Ambu, L ., Baya, W., & Knight, A. T.
(2016). Identif ying where REDD+ financially out-competes oil palm
in floodplain landscapes using a fine-scale approach. PLoS One,
11(6), 1–23 . /10.1371/journ al.pone.0156481
Agrawal, A., & Gibson, C. C. (1999). Enchantment and disenchantment:
The role of communit y in natural resource conservation. World
Development, 27(4), 629–649. htt ps://doi.or g/10.1016/S0 305 -750X
( 9 8 ) 0 0 1 6 1 - 2
Agrawal, A., & Redford, K. (2009). Place, conser vation, and displace-
ment. Conservation and Society, 7. ht tps://doi.o rg/10.4103/0972-
4923 .5 4790
Alfaro, R. I., Fa dy, B., Vendramin, G. G., Dawson, I. K., Fleming, R. A.,
Sáenz-Romero, C., Lindig-Cisneros, R. A., Murdock, T., Vinceti, B.,
Navarro, C. M., Skrøppa, T., Baldinelli, G., El-Kassaby, Y. A ., & Loo,
J. (2014). The role of forest genetic resources in responding to bi-
otic and abiotic factors in the context of anthropogenic climate
change. Forest Ecology and Management, 333, 76–87. https://doi.
Allen, A., & Chapman, D. (2001). Impacts of afforest ation on ground-
water resources and quality. Hydrogeology Journal, 9, 390–400.
htt ps:// 07/s1004 0 010 0148
Angelson, A., Brockhaus, M., Sunderlin, W. D., & Verchot, L. V. (2012).
Analysing REDD+: Challenges and choices. In A . Angelson, M.
Brockhaus, W. D. Sunderlin, & L. V. Verchot (Eds.), Analysing REDD+:
Challenges and choices. CIFOR. cif or/
Arroyo-Rodríguez, V., Melo, F. P. L., Martíne z-Ramos, M., Bongers, F.,
Chazdon, R. L., Meave, J. A ., Norden, N., Santos, B. A ., Leal, I.
R., & Tabarelli, M. (2017). Multiple successional pathways in hu-
man-modified tropical landscapes: New insights from forest suc-
cession, forest fragmentation and landscape ecology research.
Biological Reviews, 92(1), 326–3 40 . htt ps://doi. org/10 .1111/br v.
Asmelash, F., Bekele, T., & Bir hane, E. (2016). The potential role of ar-
buscular mycorrhizal fungi in the restoration of degraded lands.
Frontiers in Microbiology, 7(JUL), 1–15.
Bala, G., Caldeira, K., Wickett, M., Phillips, T. J., Lobell, D. B., Delire, C .,
& Mirin, A. (2007). Combined climate and carbon-cycle effec ts
of large-scale deforestation . Proceedings of the National Academy
of Sciences of the United States of America, 104 (16), 6550–6555.
htt ps:// s.0 60 89 9810 4
Bannister, J. R., Vargas-Gaete, R., Ovalle, J. F., Acevedo, M., Fuentes-
Ramirez, A ., Donoso, P. J., Promis, A., & Smith-Ramírez, C. (2018).
Major bottlenecks for the restoration of natural forests in Chile.
Restoration Ecology, 26(6), 1039–104 4. https: // .1111/rec.
Barlow, J., & Peres, C. A. (2008). Fire-mediated dieback and composi-
tional cascade in an Amazonian forest. Philosophical Transactions
of the Royal Society B: Biological Sciences, 363(1498), 1787–1794.
   DI SACCO et Al .
Baskin, C. C ., & Baskin, J. M. (2014). Seeds: Ecology, biogeography, and
evolution of dormancy and germination (2nd ed.). Academic Press.
Becknell, J. M., Kissing Kucek, L., & Powers , J. S. (2012). Aboveground bio-
mass in mature and secondary seasonally dry tropical forests: A lit-
erature review and global synthesis. Forest Ecology and Management,
276 , 88–95.
Bellard, C., Cassey, P., & Blackburn, T. M. (2016). Alien species as a
driver of recent extinctions. Biology Letters, 12(4 ) , ht tp s ://d o i .
Besseau, P., Graham, S ., & Chris tophersen, T. (Eds.). (2018). Restoring for-
ests and landscapes: The key to a sustainable future. Global partner-
ship on forest and landscape restoration.
Betts, R . A . (2000). Offset of the potential car bon sink from boreal for-
estation by decreases in surfa ce albedo. Nature, 408(68 09), 187
Bloomfield, G., Meli, P., Brancalion, P. H. S., Terris, E., Guariguata, M.
R., & Garen, E. (2019). Strategic insights for c apacity develop-
ment on forest landscape restoration: Implications for addressing
global commitments. Tropical Conservation Science, 12 (November).
htt ps:// /10.1177/1940 0 82919 887589
Boissière, M., Herold, M., Atmadja, S., & Sheil, D. (2017). The feasibil-
ity of local participation in Measuring, Reporting and Verification
(PMRV) for REDD+. PLoS One, 12(5), 1–9.
journ al.pone.0176897
Bond, W. J., Stevens, N., Midgley, G. F., & Lehmann, C. E. R. (2019).
The trouble with trees: Afforestation plans for Africa. Trends in
Ecology & Evolution, 34(11), 963–965.
Brancalion, P. H. S., Amazonas, N. T., Chazdon, R. L., van Melis, J.,
Rodrigues, R. R., Silva, C. C., Sorrini, T. B., & Holl, K. D. (2020).
Exotic eucalypts: From demonized trees to allies of tropical forest
restoration? Journal of Applied Ecology, 57(1), 55–66. https://doi.
org /10.1111/1365-26 64 .13513
Brancalion, P. H. S ., Bello, C., Chazdon , R. L., Galetti, M., Jordano, P., Lima,
R. A. F., Medina, A., Pizo, M. A., & Reid, J. L. (2018). Maximizing
biodiversity conservation and carbon stocking in restored tropical
forest s. Conservation Letters, 11(4), 1–9. htt ps://doi. org/10 .1111/
conl.1245 4
Brancalion, P. H. S., Cardozo, I. V., Camatta, A., Aronson , J., & Rodrigues,
R. R. (2014). Cultural ecosystem services and popular perceptions
of the benefits of an ecological restoration project in the Brazilian
Atlantic Forest. Restoration Ecology, 22(1), 65–71. https://doi.
org /10.1111/re c.120 25
Branca lion, P. H. S., & Chazdon , R. L. (2017). Beyon d hectares: Four prin-
ciples to guide refore station in the context of tropical forest and
landscape restoration. Restoration Ecology, 25(4), 491–496. https://
doi .org/10.1111/rec.12519
Brancalion, P. H. S., & Holl, K. D. (2020). Guidance for successful tree
planting initiatives. Journal of Applied Ecology, 205(4969), 2349–
2361. ht tp s://doi .o rg /10.1111 /1365-2664.1372 5
Brancalion, P. H. S., Lamb, D., Ceccon, E., Boucher, D., Herbohn, J.,
Strassburg, B., & Edwards, D. P. (2017). Using market s to leverage
investment in forest and landscape restoration in the tropic s. Forest
Policy and Economics, 85, 103–113. ht tps:// /10.1016/j.
Brancalion, P. H. S., V iani, R. A. G., Strassburg, B. B. N., & Rodrigues , R.
R. (2012). Finding money for tropical fores t restoration. Unasylva,
239(63), 41–50. 8- 014-0173-7.2
Broadhurst, L. M., Lowe, A., Coates, D. J., Cunningham, S. A ., McDonald,
M. , Vesk , P. A., & Yate s, C. (2008 ). Seed supp ly for br oa dscal e re s to -
ration: Maximizing evolutionary potential. Evolutionary Applications,
1(4), 587–597. j.1752-4571.200 8. 00 045.x
Brockerhoff, E. G., Jactel, H., Parrotta, J. A., Quine, C. P., & Sayer, J.
(2008). Plantation forests and biodiversity: Oxymoron or oppor-
tunity? Biodiversity and Conservation, 17(5), 925–951. https://doi.
org /10.1007/s1053 1-008-9380 -x
Bryan, J. E., Shearman, P. L., Asner, G. P., Knapp, D. E., Aoro, G ., & Lokes,
B. (2013). Extreme differences in forest degradation in Borneo:
Comparing practices in Sarawak, Sabah, and Brunei. PLoS One, 8(7),
e69679. htt ps:// our n al.p one .0 069679
Burrascano, S., Chytrý, M., Kuemmerle, T., Giarrizzo, E., Luyssaert,
S., Sabatini, F. M., & Blasi, C. (2016). Current European policies
are unlikely to jointly foster c arbon sequestration and protect
biodiversity. Biological Conservation, 201, 370–376. https://doi.
Burton, V., Moseley, D., Brown, C., Metzger, M. J., & Bellamy, P. (2018).
Reviewing the evidence base for the effects of woodland ex-
pansion on biodiversity and ecosystem services in the United
Kingdom . Forest Ecology and Management, 430, 366–379. https://
Calviño-Cancela, M., Rubido-Bará, M., & van Etten, E. J. B. (2012). Do
eucalypt plantations provide habitat for native forest biodiver-
sity? Forest Ecology and Management, 270, 153–162. https://doi.
Catterall, C. P. (2016). Roles of non-native species in large-scale re-
generation of moist tropical forests on anthropogenic grassland.
Biotropica, 48(6), 809–824. /10.1111/btp.12384
Chave, J., Réjou-Méchain, M., Búrquez, A., Chidumayo, E., Colgan, M. S.,
Delitti, W. B. C., Duque, A., Eid, T., Fearnside, P. M., Goodman, R.
C., Henry, M., Martínez-Yrízar, A., Mugasha, W. A ., Muller-Landau,
H. C., Mencuccini, M., Nelson, B. W., Ngomanda, A., Nogueira, E.
M., Ortiz-Malavassi, E., … Vieilledent, G. (2014). Improved allome-
tric models to estimate the aboveground biomass of tropical trees.
Global Change Biology, 20 (10), 3177–3190.
Chazdon, R. (2014). Second growth: The promise of tropical forest re-
generation in an age of deforestation. The Universit y of Chicago
Chazdon, R., & Brancalion, P. (2019). Restoring forests as a means to
many ends. An urgent need to replenish tree canopy cover calls for
holistic approaches. Science, 365(6448),
scien ce.aax9539
Chazdon, R. L., & Uriarte, M. (2016). Natural regeneration in the con-
text of large-sc ale forest and landscape restoration in t he t ropics.
Biotropica, 48(6), 709–715.
Chomba, S., Kariuki, J., Friis Lund, J., & Sinclair, F. (2016). Roots of in-
equity: How the implementation of REDD+ reinforces past injus-
tices. Land Use Policy, 50, 202–213. ht tps://
Crane, E. (2020). Woodlands for climate and nature: A review of woodland
planting and management approaches in the UK for climate change
mitigation and biodiversity conservation. Repor t to the RSPB. Royal
Societ y for the Protection of Birds. /Image
s/Fores tr y%20and %20cli mate%20cha nge%20rep or t%20Feb
Crouzeilles, R., Barros, F. S. M., Molin, P. G., Ferreira, M. S., Junqueira, A.
B., Chazdon, R . L ., Lindenmayer, D. B., Tymus, J. R . C ., Strassburg,
B. B. N., & Brancalion, P. H. S. (2019). A new approach to map land-
scape variation in forest restoration success in tropical and tem-
perate forest biomes. Journal of Applied Ecology, 56(12), 2675–2686.
https ://doi.or g/10.1111/1365 -266 4.13501
Crouzeilles, R., Beyer, H. L., Monteiro, L. M., Feltran-Barbieri, R.,
Pessôa, A. C. M., Barros, F. S. M., Lindenmayer, D. B., Lino, E. D.
S. M., Grelle, C. E. V., Chazdon, R. L., Matsumoto, M., Rosa, M.,
Latawiec, A. E., & Strassburg, B. B. N. (2020). Achieving cost-ef-
fective landscape-scale forest restoration through targeted natural
regeneration. Conservation Letters, 13(3).
conl .1270 9
Cuenca, P., Robalino, J., Arriagada, R., & Echeverria, C. (2018). Are gov-
ernment incentives effective for avoided deforestation in the
tropical Andean forest? PLoS One, 13(9), e0203545. https://doi.
org/10.1371/journ al.pone.0203545
DI SACCO et Al .
Danielsen, F., Adrian, T., Brofeldt, S., van Noordwijk, M., Poulsen, M.
K., Rahayu, S., Rutishauser, E., Theilade, I., Widayati, A., A n, N. T.,
Bang, T. N., Budiman, A., Enghoff, M., Jensen, A . E., Kurniawan, Y.,
Li, Q., Mingxu, Z., Schmidt-Vogt, D., Prixa, S., … B urgess , N. (2013).
Community monitoring for REDD+: International promises and
field realities. Ecology and Society, 18(3).
ES-05464 -180341
Dass, P., Houlton, B. Z., Wang, Y., & Warlind, D. (2018). G rasslands may be
more reliable carbon sinks than forests in California. Environmental
Research Letters, 13(7 ), 7402 7. http s://do i.o rg /10.10 88/174 8-93 26 /
de Almeida, D. R. A., Stark, S. C., Valbuena, R., Broadbent, E. N., Silva,
T. S . F., de Resende, A. F., Ferreira, M. P., Cardil, A., Silva, C. A .,
Amazonas, N., Zambrano, A. M. A., & Br ancalion, P. H. S. (2020). A
new era in forest restoration monitoring. Restoration Ecology, 28(1),
8–11. ht tp s://doi.o rg /10.1111 /re c.13067
de Souza, S. E. X. F., Vidal, E., Chagas, G. D. F., Elgar, A. T., & Brancalion, P.
H. S. (2016). Ecological outcomes and livelihood benefits of commu-
nity-managed agroforests and second growth forest s in Southeast
Brazil. Biotropica, 48(6), 868–881.
De Vitis, M., Hay, F. R., Dickie, J. B., Trivedi, C., Choi, J., & F iegener, R.
(20 20 ). S ee d st or age: Main tain in g se ed via bi lity and vigor for resto-
ration u se. Restoration Ecolo gy, 28(S3 ), 1–7. http s://doi.o rg /10.1111/
re c.13174
Deere, N. J., Guillera-arroita, G., Swinfield, T., Milodowski, D. T., Coomes,
D. A., Bernard, H., Reynolds, G., Davies, Z. G., & Struebig, M. J.
(2020). Maximizing the value of forest restoration for tropical
mammals by detecting three-dimensional habitat associations.
Proceedings of the National Academy of Sciences of the United States
of America, 117(42), 26254–26262.
20018 23117
Díaz, S., Hector, A., & Wardle, D. A. (2009). Biodiversity in forest c arbon
sequestration initiatives: Not just a side benefit. Current Opinion in
Environmental Sustainability, 1(1), 55–60. htt ps://
Díaz, S., Pascual, U., Stenseke, M ., Mar tín-López, B., Watson, R. T.,
Molnár, Z., Hill, R., Chan, K. M. A., Baste, I. A., Brauman, K. A.,
Polasky, S., Church, A., Lonsdale, M., Larigauderie, A., Leadley,
P. W., van Oudenhoven, A. P. E., van der Plaat, F., Schröter, M.,
Lavorel, S., … Shirayama, Y. (2018). Assessing nature’s contributions
to people. Science, 359(6373), 270–272.
scien ce.aap8826
Dolch, R., Ndriamiary, J., Ratolojanahary, T., Randrianasolo, M., &
Ramanantenasoa, I. (2015). Improving livelihoods, training pa-
ra-ecologists, enthralling children: Earning trust for effective
community- based biodiversity conservation in Andasibe, eastern
Madagascar. Madagascar Conservation & Development, 10(1), 21.
Douwes, E., & Buthelezi, N. (2016). G rowing forests and communities.
Journal of Geography, 29(5), 221–222.
0 0 2 2 1 3 4 3 0 0 8 9 8 7 2 9 6
Duguma, L. A., Minang, P. A., Mpanda, M., Kimaro, A., & Alemagi, D.
(2015). Landscape restoration from a social-ecologic al system
perspective? In P. A. Minang, M. van Noordwijk, O. E. Freeman, C.
Mbow, J. de Leeuw, & D. Catacut an (Eds.), Climate-smart landscapes:
Multifunctionality in practice (pp. 6373). World Agroforestry Centre
Dyde rski, M. K., & Jago dziński, A. M. (2020). Imp act of invasive tre e spe -
cies on natural regeneration species composition, diversit y, and
density. Forests, 11(4), 1–20. 0456
Elliott , S., Blakesley, D., & Hardwick, K. (2013). Restoring tropical forests.
A practical guide. Royal Botanic Gardens, Kew.
Ennos, R., Cottrell, J., Hall, J., & O'Brien, D. (2019). Is the introduction
of novel exotic forest tree species a rational response to rapid
environmental change? A British perspective. Forest Ecology and
Management, 432, 718–728.
ENSCONET. (2009a). ENSCONET Curation protocols & recommendations
(Royal Botanic Gardens, Kew (Ed.)). Royal Botanic Gardens, Kew.
ENSCONET. (20 09b). ENSCONET Seed collecting manual for wild species
(Royal Botanic Gardens, Kew & Universidad Politécnica de Madr id
(Eds.)). Royal Botanic Gardens, Kew.
Erbaugh , J. T., & Oldekop, J. A. (2018). Forest landscape restor ation
for livelihoods and well-being. Current Opinion in Environmental
Sustainability, 32, 76–83.
Erbaugh , J. T., Pradhan, N., Adams, J., Adams, J., Oldekop, J. A., Agrawal,
A., Brocking ton, D., Pritchard, R., & Chhatre, A. (2020). Global
forest restoration and the importance of prioritizing local com-
munities. Nature Ecology & Evolution., 4, 1472–1476. https://doi.
org /10.103 8/s4155 9-020 -01 282 -2
Estrada-Villegas, S., Bailón, M ., Hall, J. S., Schnit zer, S. A ., Turner, B. L.,
Caughlin, T., & van Breugel, M. (2019). Edaphic factors and initial
conditions influence successional trajectories of early regenerating
tropical dry fores ts. Journal of Ecology, 108, 160–174 . htt ps: //doi.
org /10.1111/1365-2745 .1326 3
Fagan, M. E., Reid, J. L ., Holland, M. B., Drew, J. G., & Zahawi, R.
A. (2020). How feasible are global forest restoration commit-
ments? Conservation Letters, 13(3), 1–8.
conl .1270 0
FAO. (2 019). Restoring forest landscapes through assisted natural regener-
ation (ANR) – A practical manual. Licence: CC BY-NC-SA 3.0 IGO.
Food and A griculture Organization of the United Nations, 52 p.
Feng, X., Fu, B., Piao, S., Wang, S., Ciais, P., Zeng , Z., Lü, Y., Zeng, Y.,
Li, Y., Jiang, X., & Wu, B. (2016). Revegetation in China's Loess
Plateau is approaching sustainable water resource limits. Nature
Climate Change, 6(11), 1019–1022. htt ps://doi.or g/10.1038/n clim
Florentine, S. K., Pohlman, C. L., & Westbrooke, M. E. (2016). The effec-
tiveness of different planting frameworks for recruitment of trop-
ical rainforest species on ex-rainforest land. Restoration Ecology,
24(3), 36 4–372.
Fofana, B., Sac ande, M., Blagna, F., Dibloni, T. O., Compaore, E., Sanon,
K. B., Maiga, Y., & Ouattara, A . S. (2020). Boosting land restoration
success in t he Great Green Wall through the use of symbiotic mi-
croorganisms for propagated tree seedlings. International Journal
of Biological and Chemical Sciences, 14(1), 110–125. https://doi.
Freitas, M. G ., Rodrigues, S. B., Campos-Filho, E. M., do Carmo, G. H. P., da
Veiga, J. M., Junqueira, R. G. P., & Vieira, D. L. M. (2019). Evaluating
the success of direct seeding for tropical forest restoration over
ten years . Forest Ecology and Management, 438(February), 224–232.
Friggens, N. L., Hester, A. J., Mitchell, R. J., Parker, T. C., Subke, J. A.,
& Wookey, P. A. (2020). Tree planting in organic soils does not
result in net carbon sequestration on decadal timescales. Global
Change Biology, 26(9), 5178–5188.
Gann, G. D., McDonald, T., Walder, B., Aronson, J., Nelson, C. R ., Jonson,
J., Hallett, J. G ., Eisenberg, C., Guariguata, M. R., Liu, J., Hua, F.,
Echeverría, C., Gonzales, E., Shaw, N., Decleer, K., & Dixon, K. W.
(2019). International principles and standards for the prac tice of
ecological restoration. Restoration Ecology, 27(S1), S1–S46. https://
doi .org/10.1111/rec.13 035
Gardon, F. R., de Tole do, R. M., Brentan, B. M., & Ferreira dos Santos,
R. (2020). Rainfall interception and plant communit y in young
forest restorations. Ecological Indicators, 109, 105779. ht tps://doi.
org /10.1016/j.ecoli n d. 2019.105779
Gibson, L., Lee, T. M., Koh, L. P., Bro ok, B. W., G ardner, T. A., Barlow, J.,
Peres, C. A ., Bradshaw, C. J. A., L aurance, W. F., Lovejoy, T. E., &
Sodhi, N. S. (2011). Primar y forests are irreplaceable for sustaining
   DI SACCO et Al .
tropical biodiversity. Nature, 478 (7369), 378–381. https://doi.
org /10.103 8/nat ur e10425
Gilmour, D. (2007). Applying an adaptive management approach to FLR.
Chapter 4. In J. Reitbergen-McCraken, S. Maginnis, & A. Sarre (Eds.),
The forest landscape restoration handbook (pp. 29–38). Earthscan.
Goosem , S., & Tucker, N. I. J. (2013). Repairing the rainforest (2nd ed.). Wet
Tropics Management Authorit y and Biotropic a Australia Pty Ltd.
Gunaratne, A. M. T. A., Gunatilleke, C. V. S., Gunatilleke, I. A. U. N.,
Madawala, H. M. S. P., & Bur slem, D. F. R. P. (2014). Overcoming
ecological barriers to tropical lower montane forest succession on
anthropogenic grasslands: Synthesis and future prospects. Forest
Ecology and Management, 329, 340–350.
Heilmayr, R., Echeverr ía, C., & L ambin, E. F. (2020). Impacts of Chilean
forest subsidies on forest cover, carbon and biodiversity. Nature
Sustainability, 3(9), 701–709. 3-020-
05 47- 0
Herzog, S. K., Kessler, M., & Cahill, T. M. (2002). Estimating species rich-
ness of tropical bird communities from rapid assessment data. The
Auk, 119(3), 749–769.
Hoban, S. (2019). New guidance for ex situ gene conservation: Sampling
realistic population systems and accounting for collection at-
trition. Biological Conservation, 235(May), 199–208. https://doi.
Hoban, S., & Strand, A. (2015). Ex situ seed collec tions will benefit
from considering spatial sampling design and species' reproduc-
tive biology. Biological Conservation, 187, 182–191. https://doi.
Höhl, M., Ahimbisibwe, V., Stanturf, J. A., Elsasser, P., Kleine, M., & Bolte,
A. (2020). Forest landscape restoration – What generates failure
and success? Forests, 11(9), 938.
Holl, K. D., & Br ancalion , P. H. S. (2020). Tree planting: Not a simple solu-
tion. Science, 368(6491), 580–581. ce.
Hong, S., Yin, G., Piao, S., Dybzinski, R ., Cong, N., Li, X., Wang, K.,
Peñuelas, J., Zeng, H., & Chen, A. (2020). Divergent responses of
soil organic carbon to afforestation. Nature Sustainability, 3(9), 694–
700. 3-020-0557-y
Hong, T. D., & Ellis, R. H. (1996). A protocol to determine seed storage be-
haviour. IPGRI Technical Bulletin No. 1 (J. M. M. Engels & J. Toll
(Eds.)). International Plant Genetic Resources Institute. https://doi.
or g /10 .1007/s1162 7- 017-9874 -x
Hong, T. D., Linington, S., & Ellis, R. H. (1998). Compendium of information
on seed storage behaviour. Royal Botanic Gardens, Kew.
Hopper, S. D. (2009). OCBIL theory: Towards an integrated understand-
ing of the evolution, ecolog y and conservation of biodiversity on
old, climatically buffered, infertile landscapes. Plant and Soil, 322(1),
49–86. 4-009-0068-0
Horák, J., Brestovanská, T., Mladenović, S., Kout , J., Bogusch, P., Halda, J.
P., & Zasadil, P. (2019). G reen desert? Biodiversit y patterns in forest
plantations. Forest Ecology and Management, 433, 34 3–348 . htt p s: //
Hu, J., Herbohn, J., Chazdon, R. L., Baynes, J., & Vanclay, J. (2020).
Silvicultural treatment effect s on commercial timber volume and
functional composition of a selectively logged Australian tropical
forest over 48 years . Forest Ecology and Management, 457(April),
IUCN. (2020). Guidan ce for using the IUCN glo bal standard for na ture-based
solutions. A user-friendly framework for the verification, design and
scaling up of nature-based solutions (1st ed.). IUCN. https://doi.
IUCN, & WR I. (2014). A guide to the Restoration Opportunities Assessment
Methodology (ROAM): Assessing forest landscape restoration opportu-
nities at the national or sub-national level (road-test edition). IUCN,
125 p.
Ivetić, V., & Devet aković, J. (2017). Concerns and evidence on ge-
netic diversity in planted forests. Reforesta, 3, 196. https://doi.
org/10.21750/ REFOR.3.15.39
Jalonen , R., Valette, M., Boshier, D., Duminil, J., & Thomas, E. (2018).
Forest and landscape restoration severely constrained by a lack
of attention to the quantity and quality of tree seed: Insights
from a global survey. Conservation Letters, 11(4), 1–9. https://doi.
org /10.1111/conl.12424
James, J. J., Svejcar, T. J., & Rinella, M. J. (2011). Demographic pro-
cesses limiting seedling recruitment in arid grassland restoration.
Journal of Applied Ecology, 48(4), 961–969. ht tps://doi .org /10.1111 /
James, S. H., & Coates, D. J. (20 00). Genetic systems in the sout h-west
flora: Implications for conservation strategies for Australian plant
species. Australian Journal of Botany, 48(3), 341–347. https://doi.
org /10.1071/BT99016
Kallow, S. (2014). UK National Tree Seed Project seed collecting manual.
Royal Botanic Gardens, Kew. 885/ 1
Kildisheva, O. A., Dixon, K. W., Silveira, F. A. O., Chapman, T., Di Sacco,
A., Mondoni, A., Turner, S. R., & Cross, A. T. (2020). Dormancy and
germination: Making every seed count in restoration. Restoration
Ecology, 28( S3), S2 56–S265. ht tp s://doi.o rg /10.1111 /re c.13140
Koch, J. M., & Hobbs, R. J. (2007). Syntesis: Is Alcoa successfully re-
storing a jarrah forest ecosystem after bauxite mining in Western
Australia? Res toration Ecology, 15(SUPPL . 4), 137–144. https://doi.
org/10.1111/j.1526-100X.2007.0 0290.x
Köhl, M., Neupane, P. R., & Mundhenk, P. (2020). REDD+ measure-
ment, reporting and verification – A cost trap? Implications for
financing REDD+MRV costs by result-based payment s. Ecological
Economics, 168, 106513. con.2019.
Körner, C. (2017). A matter of tree longevity. Tree longevity r ather
than growth rate controls the carbon capital of forests. Science,
355(6321), 130–131. ce.aal2449
Kull, C. A., Harimanana, S. L ., Radaniela Andrianoro, A., & Rajoelison,
L. G. (2019). Divergent perceptions of the ‘neo-Australian’ forests
of lowland eastern Madagascar: Invasions, transitions, and liveli-
hoods. Journal of Environmental Management, 229, 48–56. https:// an.2018.06.004
Lazos-Chavero, E., Zinda, J., Bennet t-Curry, A., Balvanera, P., Bloomfield,
G., Lindell, C., & Naxegr a, C. (2016). Stakeholders and tropical re-
forestation: Challenges, trade-offs, and strategies in dynamic en-
vironments. Biotropica, 48(6), 900 –914. https ://doi .or g/10.1111/
León-Lobos, P., Way, M., Aranda, P. D., & Lima-Junior, M. (2012). The
role of ex situ se ed banks in the conservation of plant diversit y and
in ecological restoration in Latin Amer ica. Plant Ecolog y & Diversity,
5(2), 245–258. 874.2012.713402
Lewis, S. L., Wheeler, C. E., Mitchard, E. T. A., & Koch, A. (2019).
Regenerate natural fores ts to store carbon. Nature, 568, 25–28.
Liu, Q., Zhang, Q., Yan, Y., Zhang, X., Niu, J., & Svenning, J.-C. (2020).
Ecological restoration is the dominant driver of the recent re-
versal of desertification in the Mu Us Desert (China). Journal of
Cleaner Production, 268, 122241. ro.
Ludwig, J. A., & Reynolds, J. F. (1988). Statistical e cology: A prime r on meth-
ods and computing. Wi le y.
Luyssaert, S., Schulze, E. D., Börner, A., Knohl, A., Hessenmöller, D., Law,
B. E., Ciais, P., & Grace, J. (2008). Old-growth forests as global car-
bon sinks. Nature, 455, 213–215.
e0 7276
Madsen, M. D., Kostka, S. J., Inouye, A. L., & Zvirzdin, D. L . (2012).
Postfire restoration of soil hydrology and wildland vegetation
using sur factant seed coating technology. Rangeland Ecology and
Management, 65(3), 253–259. https ://doi .or g/10. 2111/REM -D -11-
DI SACCO et Al .
Martin, A. R., & Thomas, S. C. (2011). A reassessment of carbon content
in tropical trees. PLoS One, 6(8), e23533.
journ al.pone.0023533
Mattana, E., Peguero, B., Di Sacco, A., Agr amonte, W., Encarnación
Castillo, W. R., Jiménez, F., Clase, T., Pritchard, H. W., Gómez-
Barreiro, P., Castillo-Lorenzo, E., Terrero Encarnación, M., Way,
M. J., García, R., & Ulian, T. (2020). Assessing seed desiccation re-
sponses of native trees in the Caribbean. New Forests, 51(4), 705–
721. https://doi. org /10.10 07/s1105 6 -019- 09753 -6
Maxwell, S. L., Evans, T., Watson, J. E. M., Morel, A., Grantham, H.,
Duncan, A., Harris, N., Potapov, P., Runting, R. K., Venter, O., Wang,
S., & Malhi, Y. (2019). Degradation and forgone removals increase
the carbon impac t of intac t forest loss by 626%. Science Advances,
5(10), eaax2546.
McAlpine, C., C atterall, C. P., Nally, R . M., Lindenmayer, D., Reid, J. L .,
Holl, K. D., Bennett, A. F., Runting, R. K., Wilson, K., Hobbs, R. J.,
Seabrook, L., Cunningham, S., Moilanen, A., Maron, M., Shoo, L.,
Lunt, I., Vesk, P., Rumpff, L., Martin, T. G., … Possingham, H. (2016).
Integrating plant- and animal-based perspectives for more effective
restoration of biodiversity. Frontier s in Ecolog y and the Environment,
14(1), 37–45.
Meli, P., Holl, K . D., Benayas, J. M. R., Jones, H. P., Jones, P. C., Montoya,
D., & Mateos, D. M. (2017). A global review of past land use, cli-
mate, and active vs. passive restoration effects on forest recov-
er y. PLoS One, 12(2), e0171368. al.
Merrit t, D. J., & Dixon, K . W. (2011). Restoration seed banks – A mat-
ter of scale. Science, 332(6028), 424–425. /10.1126/
scien ce.1203083
Meyfroidt, P., Rudel, T. K ., & Lambin, E. F. (2010). Forest transitions,
trade, and the global displacement of land use. Proceedings of the
National A cademy of Science s of the United States of Ame rica, 107(49),
20917–2092 2. https://doi. org /10.1073/pnas.10147 73107
Millar, C. I., & Stephenson, N. L. (2015). Temperate forest health in an
era of emerging megadisturbance. Science, 349(6250), 823–826. ce.aaa9933.
Molin, P. G., Chazdon, R ., de Barros, F., Ferraz, S., & Brancalion, P. H. S.
(2018). A landscape approach for cost-effective large-sc ale fores t
restoration. Journal of Applied Ecology, 55(6), 2767–2778. https://
doi .org/10.1111/1365 -266 4.1326 3
Moomaw, W. R., Masino, S. A., & Faison, E. K. (2019). Intact forests in the
United States: Proforest ation mitigates climate change and ser ves
the greatest good. Frontiers in Forests and Global Change, 2(June),
NCC (Natural Capital Committee). (2020). Advice on using nature
based inter ventions to reach net zero greenhouse gas emissions by
2050. Department for Environment, Food and Rural Affairs, HM
Government, 26 p.
Nepstad, D., Mc Grath, D., Stickler, C ., Alencar, A., Azevedo, A., Swet te,
B., Bezerra, T., DiGiano, M., Shimada, J., Da Motta, R. S., Armijo, E.,
Castello, L., Brando, P., Hansen, M. C., McGrath-Horn, M., Carvalho,
O., & Hess, L. (2014). Slowing Amazon deforestation through pub-
lic policy and inter ventions in beef and soy supply chains. Science,
344(6188), 1118–1123. ce.1248525
Neuenkamp, L., Prober, S. M., Price, J. N., Zobel, M., & Standish, R. J.
(2019). Benefits of mycorrhizal inoculation to ecological resto-
ration depend on plant functional type, restoration context and
time. Fungal Ecology, 40, 140–149.
Newmark, W. D., Jenkins, C. N., Pimm, S. L., McNeally, P. B., & Halley,
J. M. (2017). Targeted habitat restoration can reduce extinction
rates in fragmented fores ts. Proceedings of the National Academy
of Sciences of the United States of America, 114(36), 9635–9640. 34114
Noormets, A., Epron, D., Domec, J. C., McNulty, S. G ., Fox, T., Sun, G.,
& King, J. S. (2015). Ef fect s of forest management on produc tivit y
and carbon sequestration: A review and hypothesis. Forest Ecology
and Management, 355, 124–140.
NYDF Assessment Partners. (2019). Protecting and restoring forests: A
story of large commitments yet limited progress. New York declaration
on forests five-year assessment report. Climate focus (coordinator and
Oldekop, J. A., Sims, K. R. E., Karna, B. K., Whittingham, M. J., & Agrawal,
A. (2019). Reductions in defo res tat ion and povert y from decentr al-
ized forest management in Nepal. Nature Sustainability, 2(5 ), 42 1–
428. 3-019-0277-3
Ong, C. K., Black, C. R., & Muthuri, C. W. (2006). Modif ying forestr y and
agroforestry to increase water productivity in the semi-arid tropics.
CAB Revie ws: Perspectives i n Agriculture, Veter inary Science , Nutrition
and Natural Resources, 1(065). NR
200 61065
Parsa, V. A., Salehi, E., Yavari, A. R., & van Bodegom, P. M. (2019).
Evaluating the potential contribution of urban ecosystem service
to climate change mitigation. Urban Ecosystems, 22(5), 989–1006. 2-019-00870 -w
Pedrini, S., & Dixon, K. W. (2020). International principles and standards
for native seeds in ecologic al restoration. Restoration Ecolog y,
28(S 3), S286–S3 03. htt ps://doi. org/10 .1111/rec.13155
Pedrini, S., Dixon, K. W., & Kildisheva, O. A. (2020). Seed enhance-
ment: Getting seeds restoration-ready. Restoration Ecology, 28(S3).
https ://doi.or g/10.1111/r ec .13184
Pedrini, S., Gibson-Roy, P., Trivedi, C., Gálvez-Ramírez, C., Hardwick, K .,
Shaw, N., Frischie, S., Laverack, G., & Dixon , K. (2020). Collection
and produc tion of native seeds for ecological restoration.
Restoration Ecology, 28(S3), S228–S2 38 . ht tp s://doi.o rg /10.1111/
rec .13190
Perino, A ., Pereir a, H. M., Navarro, L. M., Fernández, N., Bullock, J.
M., Ceausu, S., Cortés-Avizanda, A., van Klink , R., Kuemmerle, T.,
Lomba, A., Pe'er, G., Plieninger, T., Rey Benayas, J. M., Sandom, C .
J., Svenning, J., & Wheeler, H. C. (2019). Rewilding complex eco-
systems. Science, 364, eaav5570.
ce.a av5570
Pfund, J.-L., & Robinson, P. (Eds.). (2005). Non-timber forest products:
Between poverty alleviation and market forces. Intercooperation,
50 p.
Philipson, C. D., Cutler, M. E. J., Brodrick, P. G ., Asner, G . P., Boyd, D.
S., Moura Costa, P., Fiddes, J., Foody, G. M., van der Heijden, G.
M. F., Ledo, A., Lincoln, P. R., Margrove, J. A ., Martin, R. E., Milne,
S., Pinard, M. A., Reynolds, G., Snoep, M., Tangki, H., Sau Wai, Y.,
… Burslem, D. F. R. P. (2020). Active restoration accelerates the
carbon recovery of human-modified tropical forests. Science,
369(6505), 838–841. ce.aay4490
Poorter, L ., Bongers, F., Aide, T. M., Almeyda Zambrano, A. M.,
Balvanera, P., Becknell, J. M., Boukili, V., Brancalion , P. H. S.,
Broadbent, E. N., Chazdon, R. L., Craven, D., de Almeida- Cortez,
J. S., Cabral, G. A . L., de Jong, B. H. J., Denslow, J. S., Dent, D.
H., DeWalt, S. J., Dupuy, J. M., Durán, S. M., … Rozendaal, D.
M. A. (2016). Biomass resilience of Neotropical secondar y for-
ests. Nature, 530(7589), 211–214. 38/natur
Porras, I., Grieg-Gran, M., & Neves, N. (2008). All that glitters: A review
of payments for watershed services in developing countries. Natural
Resource Issues (No. 11). International Institute for Environment
and Development , 130 p. ISBN: 978-1-84369-653-7.
Reid, J. L., Fagan, M. E., Lucas, J., Slaughter, J., & Z ahawi, R. A . (2019).
The ephemeralit y of secondary fores ts in southern Cost a Rica.
Conservation Letters, 12(2), 1–7.
Rey Benayas, J. M., New ton, A. C., Diaz, A., & Bullock, J. M. (20 09).
Enhancement of biodiversity and ecosystem services by ecologi-
cal restoration: A meta-analysis. Science, 325(5944 ), 1121–1124. ce.1172460
   DI SACCO et Al .
Rezende, G. M., & Vieira, D. L. M. (2019). Forest restoration in southern
Amazonia: Soil preparation triggers natural regeneration. Forest
Ecology and Management, 433, 93–104. https: //doi.or g/10.1016/j.
Richardson, D. M., & Kluge, R. L . (20 08). Seed banks of invasive
Au str alian Ac a ci a spec ies in So ut h Afr ica : Role in in va siven e ss an d
options for management. Perspectives in Plant Ecology, Evolution
and Systematics, 10(3), 161–177.
Ricketts, T. H., Daily, G. C., Ehrlich, P. R., & Michener, C. D. (2004).
Economic value of tropical forest to coffee production. Proceedings
of the National Academy of Sciences of the United States of America,
101(34), 12579–12582. /10.1073/pnas.04051 47101
Ritter, C. D., Häg gqvist, S., Karlsson , D., Sääksjärvi, I. E., Muasya, A . M.,
Nilsson, R. H., & Antonelli, A. (2019). Biodiversity assessments in
the 21st century : The potential of insect traps to complement en-
vironmental samples for estimating eukar yotic and prokaryotic di-
versity using high-throughput DNA metabarcoding. Genome, 62(3),
Rozendaal, D. M. A., Bongers, F., Aide, T. M., Alvarez-Dávila, E., Ascarrunz,
N., Balvanera, P., Becknell, J. M., Bentos, T. V., Brancalion, P. H.
S., Cabral, G. A. L., Calvo-Rodriguez, S., Chave, J., César, R. G.,
Chazdon, R. L., Condit, R., Dallinga, J. S., De Almeida-Cortez, J. S.,
De Jong, B., & De Oliveira, A., … Poorter, L. (2019). Biodiversity
recover y of Neotropical secondar y forests. Science Advances, 5(3),
eaa u3114 . ht tps:// /10.1126/s ciadv. aa u3114
Scott, D. F., & Prinsloo, F. W. (2008). Longer-term ef fec ts of pine and eu-
calypt plantations on streamflow. Water Resources Research, 45(7),
1–8. R006781
Seddon , N., Chausson, A ., Berr y, P., Girardin, C. A . J., Smith, A., & Turner,
B. (2020 ). Unde rs ta ndi ng t he va lue and limits of natur e-b ased so lu-
tions to climate change and other global challenges. Philosophical
Transactions B: Biological Sciences, 375. ht tps://doi.o rg/10.1098/
Seddon , N., Turner, B., Berry, P., Chausson, A ., & Girardin, C . A. (2019).
Grounding nature-based climate solutions in sound biodiversity
science. Nature Climate Change, 9, 84–87.
s4155 8-019-0405-0
Seymour, F., & Busch, J. (2016). Why forests? Why now? The science, eco-
nomics, and politics of tropical forests and climate change. Center for
Global D evelopment. ISBN: 978-1-933286-85-3.
Shaw, N., Barak, R. S., Campbell, R. E., Kirmer, A., Pedrini, S., Dixon, K., &
Frischie, S. (2020). Seed use in the field: Delivering see ds for resto-
ration su ccess. Restoratio n Ecology, 28(S3).
rec .13210
Shono, K ., Cadaweng, E. A ., & Durst, P. B. (2007). Application of assisted
natural regeneration to restore degraded tropical forestlands.
Restoration Ecology, 15(4), 620–626. http s://doi.o rg /10.1111 /j.1526 -
Soares-Filho, B., Rajão, R., Macedo, M., Carneiro, A., Costa, W., Coe,
M., Rodrigues, H., Alencar, A. C., & Code, B. F. (2014). Cracking
Brazil's Forest Code. Science, 344(6182), 363–364. https://doi.
org/10.1126/scien ce.1246663
Steidinger, B. S., Crowther, T. W., Liang, J., Van Nuland, M. E., Werner,
G. D. A., Reich, P. B., Nabuurs, G., De-Miguel, S., Zhou, M., Picard,
N., Herault, B., Zhao, X., Zhang, C., Routh, D., Peay, K. G ., Abeg g,
M., Adou Yao, C. Y., Alber ti, G., Almeyda Zambrano, A., … Zo-Bi, I.
C. (2019). Climatic controls of decomposition drive the global bio-
geography of forest-tree symbioses. Nature, 569(7756), 404–408. 6-019-1128-0
Steur, G ., Verburg, R. W., Wassen, M. J., & Verweij, P. A . (2020).
Shedding light on relationships between plant diversity and trop-
ical forest ecosys tem services across spatial scales and plot sizes.
Ecosystem Services, 43(March), 101107. ht tps://doi.o rg/10.1016/j.
Stewart, M., & Blomley, T. (1994). Use of Melia volkensii in a semi-arid
agroforestry system in Kenya. Commonwealth Forestry Review,
73(2), 128–131.
TEEB. (2009). TEEB climate issues update (Issue September). Retrieved
from http://www.teebw catio n/clima te-issue s-up
dat e/
ter Steege, H., Prado, P. I., Lima, R . A . F. D., Pos, E., de Souza Coelho, L.,
de Andrade Lima Filho, D., Salomão, R. P., Amaral, I. L., de Almeida
Matos, F. D., C astilho, C . V., Phillips, O. L., Guevara, J. E., de Jesus
Veiga Carim, M., Cárdenas López, D., Magnusson, W. E., Wittmann,
F., Ma rtins , M. P., Sa ba tie r, D., Ir um e, M. V., … Pic kavan ce , G. (202 0) .
Biased-corrected richness estimates for the Amazonian tree flora.
Scientific Reports, 10(1) , 1–13. htt ps://doi.or g/10.1038/s 4159 8-
020-66686 -3
Thomas, E., Jalonen, R., Loo, J., Boshier, D., Gallo, L ., Cavers, S., Bordács,
S., Smith, P., & Bozzano, M. (2014). Genetic considerations in
ecosystem restoration using native tree species. Forest Ecology
and Management, 333(2014), 66–75. ht tps://doi.o rg/10.1016/j.
Tucker, N. I. J., & Simmons, T. (20 09). Restoring a rainfores t habitat linkage
in north Queensland: Donaghy’s Corridor. Ecological Management
and Restoration, 10(2 ), 98–112. ht tps://doi .org/10.1111/j.14 42-
UN Convention on Biological Diversity. (2011). Nagoya protocol on ac-
cess to genetic resources and the fair and equitable sharing of benefits
arising from their utilization to the convention on biological diver sity.
Se cr eta riat of th e Conv en tio n on Biol ogi ca l Dive rsi t y. ht tps://w w w. col/nagoy a-proto col-en.pdf
UNFCCC. (2008). Report of the conference of the parties on its thir teenth
session, held in Bali from 3 to 15 December 2007. United Nations
Framework Convention on Climate Change.
UNFCCC. (2011). Report of the conference of the par ties on it s sixteenth
session, held in Cancun from 29 November to 10 December 2010.
United Nations Framework Convention on Climate Change.
Urzedo, D. I., Vidal, E., Sills, E. O., Pinã-Rodrigues, F. C. M., & Junqueira,
R. G. P. (2016). Tropical forest seeds in the household economy:
Effects of market participation among three sociocultural groups
in the Upper Xingu region of the Brazilian Amazon. Environmental
Conservation, 43(1), 13–23 . https: //doi.o rg /10.1017/S0376 89291
Veldman, J. W., Aleman, J. C., Alvar ado, S. T., Anderson, T. M., Archibald,
S., Bond, W. J., Boutton, T. W., Buchmann, N., Buisson, E., Canadell,
J. G ., Dechoum, M. S., Diaz-Toribio, M. H., Durigan, G., Ewel, J. J.,
Fernandes, G. W., Fidelis, A ., Fleischman, F., Good, S. P., & Grif fith,
D. M., … Zaloumis, N. P. (2019). Comment on “The global tree
restoration potential”. Science, 366(6463), eaay7976. https://doi.
org/10.1126/scien ce.aay7976
Veldman, J. W., Overbeck, G. E., Negreiros, D., Mahy, G., Le Stradic, S.,
Fernandes, G. W., Durigan, G., Buisson, E., Put z, F. E., & Bond, W. J.
(2015). Where tree planting and forest expansion are bad for bio-
diversity and ecosystem services. BioScience, 65(10), 1011–1018. i/biv118
Verheyen, K., Vanhellemont , M., Auge, H., Baeten, L., Baraloto,
C., Barsoum, N., Bilodeau-Gauthier, S., Bruelheide, H.,
Castagneyrol, B., Godbold, D., Haase, J., Hector, A ., Jactel, H.,
Kori che va, J. , Lor eau , M ., Mer eu, S. , M es sier, C ., Mu ys, B. , N ole t,
P., … Scherer-Lorenzen, M. (2016). Contributions of a global net-
work of tree diversity experiments to sustainable forest plan-
tations. Ambio, 45(1), 29–41. https://doi.o rg /10.10 07/s1328 0-
Wangpakapattanawong, P., Kavinchan, N., Vaidhayakarn, C., Schmidt-
Vogt, D., & Elliott , S. (2010). Fallow to forest: Applying indigenous
and scientific knowledge of swidden cultivation to tropical forest
restoration. Forest Ecology and Management, 260(8), 1399–1406.
DI SACCO et Al .
Watson, J. E. M., Evans, T., Venter, O., W illiams, B., Tulloch, A., Stewar t,
C., Thompson, I., Ray, J. C., Murray, K., Salazar, A ., McAlpine, C.,
Potapov, P., Walston, J., Robinson, J. G., Painter, M., Wilkie, D.,
Filardi, C., Laurance, W. F., Houghton, R. A., … Lindenmayer, D.
(2018). The exceptional value of intac t forest ecosys tems. Nature
Ecology & Evolution, 2(4), 599–610. https://doi.or g/10.1038/s4155
Wheeler, C. E., O meja, P. A., Chapman, C. A ., Glipin, M., Tumwesigye,
C., & Lewis, S. L. (2016). Carbon sequestration and biodiversity
following 18 years of active tropical forest restoration. Forest
Ecology and Management, 373, 44–55.
Whit te t, R. , Cott re ll , J ., Cav er s, S., Pec ur ul, M., & Enn os , R. (20 16) . Su pp ly ing
trees in an era of environmental uncertainty: Identifying challenges
faced by the forest nursery sec tor in Great Brit ain. Land Use Policy,
58, 415–426. sepol.2016.07.027
Williams, M. I., Dumroese, R. K., Page-Dumroese, D. S., & Hardegree,
S. P. (2016). Can biochar be used as a seed coating to improve na-
tive plant germination and growth in arid conditions? Journal of Arid
Environments, 125(February), 8–15.
Wyse, S. V., & Dickie, J. B. (2017). Predicting the global incidence of
seed desiccation sensitivity. Journal of Ecology, 105 (4), 1082–1093.
https ://doi.or g/10.1111/1365 -2745.12725
Wyse, S. V., & Dickie, J. B. (2018). Taxonomic affinity, habitat and seed
mass strongly predict seed desiccation response: A boosted re-
gression trees analysis based on 17 539 species. Annals of Botany,
121(1), 71–83.
Zahawi, R. A., Holl, K. D., Cole, R. J., & Leighton Reid , J. (2013). Testing
applied nucleation as a strategy to facilitate tropical forest re-
cov er y. Journal of Applied Ecology, 50(1), 88–96. https://doi.
org /10.1111/1365-26 64 .12014
How to cite this article: Di Sacco A , Hardwick KA, Blakesley
D, et al. Ten golden rules for reforestation to optimize carbon
sequestration, biodiversity recovery and livelihood benefits.
Glob Change Biol. 2021;00 :1–21. ht tp s://doi.o rg /10.1111/
... Global spatial prioritization provides initial guidelines, but for execution and greater effectiveness, an assessment must be made at the landscape or lower cartographic scale, involving historical, ecological, and socioeconomic factors [24]. A multifunctional landscape approach that improves the coexistence of different land uses with the interests of the stakeholders involved, together with a relevant review of land-cover maps and statistical analysis, is essential [25]. ...
... Finding a transdisciplinary approach that addresses the complexity of ecosystems and socio-ecological systems, considering ecological vitality, economic potential, and social acceptance, could accelerate restoration efforts, obtaining collateral benefits for all stakeholders involved [26,27]. Communication must be clear and transparent among parties, based on fair governance and incentivized by a long-term mechanism [24], with persistent processes over time [28]. Now, by understanding the perception of the importance of landscape restoration processes and the possible location of these activities on their properties, communities can provide guidelines for approaching future landscape restoration processes. ...
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Landscape restoration activities must be conducted through a transdisciplinary process, integrating social, economic, environmental, and governance aspects. Combining visions from the natural and social sciences is a challenge in highly complex territories, where unique ecosystem characteristics, economic processes, stakeholders of diverse nature, and different normativity converge. The harmonization of multiple techniques, such as multicriteria spatial analysis, expert knowledge elicitation, and social mapping, allows for an approach to defining landscape restoration areas in complex regions. This paper employs multiple techniques to define ecosystem restoration areas in a complex Colombian Andes landscape, integrating ecological and social components for sustainable development. We observed that areas of high and very high feasibility for ecological restoration, encompassing 179.5 hectares (4.84% of the study area), are predominantly located near primary forests. Although some areas have a low feasibility for conservation processes, they should not be disregarded as they still require protection. Landowners prioritize watershed and soil restoration as the most important landscape restoration activity due to their interest in improving water-related ecosystem services. This proposal enables the identification of areas with a higher restoration potential at the property level, facilitating prioritization and investment allocation for future implementation.
... According to a Bloomberg report, in 2013, India intended to offer electricity at a reasonable cost with this ultra-mega solar power plant proposal. For large projects, solar power/electricity prices have been as low as 4.34 cents per kWh [32]. Between 2012 and 2016, it was shown that the average trading of these solar systems increased by 47% each year, supplying electricity to almost 90,000 houses [33]. ...
... India's energy departments, such as MNRE, IREDA (India Renewable Energy Development Agency), and SEC (Solar Energy Center) are actively supporting and promoting projects on solar power. In February 2016, the government of capital state Delhi declared the incentives for solar power generation; as per the scheme government will pay 2 Rs./ kWh of solar power produced to charitable organizations and domestic consumers [32]. This incentive was released for two years to escalate the adoption of solar energy. ...
In Paris Climate Agreement (COP 21) in December 2015, India committed to cut its carbon (C) emissions by 30-35% of its GDP (gross domestic product) and attain 40% of installed energy from renewables by 2030. In the Glasgow Climate Pact (COP 26), an agreement developed at COP26 of UNFCCC in November 2021, India committed to enhancing its renewable energy (RE) capacity to 500 GW and installing 50% of its power from renewable resources by 2030. It is projected that this will help India to reduce its C emissions by 1Bt, reduce emission intensity to the economy by 45% by 2030, and meet the goal of C neutral economy by 2070. Realization of these highly ambitious commitments made at COP26 can be determined by the nation's growth in REs after COP21. Therefore, this article reviews the overall development of renewable energies-solar, wind, biomass, hydrogen, and hydro-in India after COP21 by the end of December 2021. This review provides detailed information on various policies and roadmaps for each energy sector commitments. It identifies major technical and social challenges associated with the sustainability of these sectors. This review offers insights into how realistically ambitious plans India has for 'phasing down' the use of coal by 2050 and being a carbon-neutral economy by 2070. India achieved massive success through solar energy after COP21 and 80% of future investments are for solar energy. Comparatively, there has been the least investment in other REs such as bio-energy, water, and wind.
... Secondly, these indicators do not measure ecological functionality [45,25] and how forests and species within them provide ecosystem services, especially at the landscape scale. Several recent articles have criticized this lack of rigor concerning ecological functionality or forest quality in FLR and associated forest restoration efforts [46][47][48][49]. Thirdly, these metrics omit entirely the human dimension intrinsic to FLR. ...
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Purpose of Review Forest landscape restoration (FLR) is an approach to restoring forest ecosystems first defined in 2000 that has gained prominence since the launch of the Bonn Challenge in 2011. FLR aims to enhance ecological integrity and improve human well-being within (forested) landscapes. The monitoring of FLR is essential to ensure effective implementation and to learn from practice. Yet to this day, monitoring remains a major challenge for FLR. Monitoring FLR requires measures at a landscape scale and across social and natural sciences. We explore some of the monitoring challenges raised by these dimensions of FLR. We assess the current theory and practice behind FLR monitoring and how it relates to practices in related environmental disciplines. Recent Findings We highlight the challenges raised by the recent attempts at monitoring FLR and explore lessons from other related fields and conclude by proposing a framework of the basic issues to consider when monitoring FLR. Summary
... In addition, economic variables have always been an element that cannot be avoided in the study of forest carbon sequestration issues, and there is controversy over whether they are effective and to what extent. In the study of forest carbon sequestration projects in Mozambique, scholars found that farmers' income from carbon sequestration projects did not produce significant carbon sequestration effects [44,45]; other scholars believe that income from carbon sequestration projects can have a positive effect on carbon sequestration, but the visibility and sustainability of income need to be ensured [46]. Therefore, this article believes that forestry income, especially the differences in forestry income between regions (A2a), should be taken int