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The role of palaeoecology in
reconciling biodiversity
conservation, livelihoods and
carbon storage in Madagascar
Lindsey Gillson
1
*, Estelle Razanatsoa
1
,
Andriantsilavo Hery Isandratana Razafimanantsoa
1
,
Malika Virah-Sawmy
2
and Anneli Ekblom
3
1
Plant Conservation Unit, Department of Biological Sciences, University of Cape Town,
Rondebosch, South Africa,
2
Sensemakers Collective, Berlin, Germany,
3
Department of Archaeology
and Ancient History, University of Uppsala, Uppsala, Sweden
Planting trees is proposed as an important climate mitigation tool, but can be
detrimental to biodiversity and livelihoods if not carefully planned and managed,
with landscape history and livelihoods in mind. In Madagascar, deforestation is of
concern, and a threat to forest-adapted biota. However, much of Madagascar’s
landscape harbours ancient mosaic and open ecosystems that are home to
unique suites of flora and fauna and provide a wide range of ecosystem services.
Though guidelines for ecologically and socially responsible reforestation are
emerging, the potential role of landscape history and palaeoecology has been
generally underemphasised. Here, using Madagascar as a case study, we argue
that forest restoration projects need a sound understanding of landscape history
that includes a greater integration of palaeoecological data. This would help
establish the former composition and extent of forests and also investigate the
antiquity of open and mosaic ecosystems. When economic interests are strong,
information from palaeoecology and environmental history can help reduce
biases when identifying appropriate locations and suites of species for
forestation. Furthermore, a reflective approach to landscape history can
contribute to restoration projects that integrate cultural and livelihood
considerations. A transdisciplinary approach that considers local needs and
cultural context can facilitate the design and implementation of restoration
projects that share benefits equitably. Underpinning this ambition is a more
comprehensive consideration of ecosystem service benefits in a changing
climate that includes accurate carbon storage calculations, as well as other
ecosystem services including water provision, soil formation and erosion
prevention, grazing resources, medicine and cultural components.
KEYWORDS
ecosystem services, livelihoods, palaeoecology, reforestation, restoration
Frontiers in Conservation Science frontiersin.org01
OPEN ACCESS
EDITED BY
Maria M. Romeiras,
University of Lisbon, Portugal
REVIEWED BY
Matthew E. Aiello-Lammens,
Pace University, United States
Dirk Hölscher,
University of Göttingen, Germany
Bruno Salomon Ramamonjisoa,
University of Antananarivo, Madagascar
*CORRESPONDENCE
Lindsey Gillson
lindsey.gillson@uct.ac.za
RECEIVED 31 August 2023
ACCEPTED 24 November 2023
PUBLISHED 20 December 2023
CITATION
Gillson L, Razanatsoa E,
Razafimanantsoa AHI, Virah-Sawmy M and
Ekblom A (2023) The role of
palaeoecology in reconciling
biodiversity conservation, livelihoods
and carbon storage in Madagascar.
Front. Conserv. Sci. 4:1286459.
doi: 10.3389/fcosc.2023.1286459
COPYRIGHT
© 2023 Gillson, Razanatsoa,
Razafimanantsoa, Virah-Sawmy and Ekblom.
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under the terms of the Creative Commons
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distribution or reproduction in other
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author(s) and the copyright owner(s) are
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which does not comply with these terms.
TYPE Review
PUBLISHED 20 December 2023
DOI 10.3389/fcosc.2023.1286459
1 Introduction
Madagascar is a biodiversity hotspot (Myers et al., 2000;
Antonelli et al., 2022) and a focus of conservation effort for
several decades (Waeber et al., 2016). The island is one of the
countries that have received the most external carbon mitigation
and afforestation funding (Neudert et al., 2017). Yet, assessment of
the last decades shows a discouraging lack of success in terms of
both biodiversity protection and local livelihoods (Tsayem Demaze,
2014;Neudert et al., 2017). Analysis of historical national tree cover
maps and satellite imagery show deforestation and forest
fragmentation in several areas of Madagascar since the 1950s,
with an increasing rate since 2005 (Vieilledent et al., 2018). In
March 2019, the island’s new presidency announced a reforestation
plan at the One Planet summit in Nairobi, committing to reforest at
least 40,000 hectares per year for the next five years (Mandimbisoa,
2021). The goal is to restore 4 million ha of “degraded forests”in the
island by 2030 within the northwest, east and central highlands
regions (Lacroix et al., 2016;Ranjatson et al., 2019).
While reforestation of degraded forest areas can have
biodiversity and carbon benefits if carefully managed, such
initiatives will also have negative consequences for both people
and biodiversity if they extend beyond formerly forested areas, or if
inappropriate species are chosen (Bond et al., 2019;Vetter, 2020).
Inadequate restoration may result when there are large economic
interests at stake, for example for carbon or mining projects.
Furthermore, the benefits of tree-planting risks being over-
estimated if only above-ground biomass is considered, as most
carbon is below ground (e.g. see Dass et al., 2018;Razafindrakoto
et al., 2018;Veldman et al., 2019). In recent years, guidelines have
emerged that advocate more ecologically sound and socially
inclusive approaches to tree-planting, and that consider ecological
relevance, drivers of forest loss, and social dimensions (e.g. see
Brancalion and Holl, 2020;Di Sacco et al., 2021). However, though
several such guidelines highlight the importance of distinguishing
reforestation of degraded forests from the afforestation of
landscapes in areas not formerly forested (Veldman et al., 2015;
Boissière et al., 2021;Di Sacco et al., 2021), there has been little
attempt to address how that distinction is established. Where
afforestation is defined, it usually refers to tree plantations on
land that did not harbor forest in the past 50 years (Bredemeier
and Dohrenbusch, 2009). Yet, this baseline in the 1970s is unlikely
to be an accurate reflection of the long-term history and ecological
character of landscapes. Many landscapes have been radically
transformed at least since the “great acceleration”of human
impact in the 1950s, and most likely since the dramatic changes
of the C18
th
and C19
th
including, industrialization and agricultural
intensification, and European colonization (Dietl et al., 2015;Barak
et al., 2016;Falk et al., 2019;Gillson et al., 2021).
Tree planting in areas that were not historically forested may
negatively impact biodiversity and ecosystem services provided by
ancient open and mosaic landscapes (Bond et al., 2019;Vetter, 2020;
Garcıa-Dory et al., 2021;Lehmann et al., 2021). Afforestation
projects commonly use exotic species and compromise other
ecosystem services including water provision, grazing/forage, and
biodiversity (Stickler et al., 2009;Bond et al., 2019;Garcı
a-Dory
et al., 2021). Loss of access to such ecosystem services is of particular
concern given that Madagascar is one of the world’s poorest
countries, with almost 80% living below the poverty line $1.90 a
day (Neugarten et al., 2016;Baumann, 2021). Most rural
communities in Madagascar are directly dependent on ecosystem
services, and multiple livelihood streams are essential in buffering
environmental shocks (Huff, 2014). While carbon markets are
frequently touted as a win-win solution for both climate and
people, payments for ecosystem services may not reach the most
vulnerable and can furthermore create a dependence on short-term
external payments that are not sustainable and would erode social-
ecological resilience and adaptive capacity in the longer-term (Bulte
et al., 2008;Gross-Camp et al., 2012;Kronenberg and Hubacek,
2013). It is therefore vital that attempts to mitigate climate change
consider local livelihoods and biodiversity and integrate more
accurate carbon storage calculations alongside consideration of
other ecosystem services.
Here, we advocate for the integration of palaeoecological
approaches that can be used to explore former forest extent and
history over timescales that are ecologically relevant, using case
studies from Madagascar (Virah-Sawmy, 2009;Razafimanantsoa,
2022). Such palaeoecological methods include the analysis of fossil
pollen from lake sediments, the analysis of charcoal to reconstruct
fire history, stable isotopes to investigate vegetation and climate,
dung fungal spores to reconstruct changes in the abundance of
herbivores, diatoms to examine water quality, and geochemistry to
investigate the interacting effects of climate and disturbance. We
argue that determining appropriate reference conditions for
restoration requires establishing the extent and composition of
vegetation over time-periods that are relevant to landscape
history and allow the impact of environmental change to be
assessed over timescales of decades to centuries and even
thousands of years. In Madagascar, human settlement (around
2000 years ago, Douglass et al., 2019), the Medieval Climate
Anomaly (around 1000 years ago) and European colonization
since the 19
th
Century, (Rid, 2010) provide critical reference
conditions, that can help to distinguish ancient open and mosaic
ecosystems from formerly forested areas. Research around such
milestones in history, also contributes to disentangling
anthropogenic and environmental drivers of land-cover change.
This information is valuable in exploring the resilience of social-
ecological systems, providing an underpinning for explorations of
future sustainability under different scenarios of climate and land-
use change.
2 The importance of landscape
history–where and how should
reforestation take place
There is no doubt that deforestation has taken place in some
areas of Madagascar particularly in the last 50 years (Zaehringer
et al., 2015;Vieilledent et al., 2018;Suzzi-Simmons, 2023).
However, the idea that the whole island was once entirely covered
by forest has been disproved by a growing number of
Gillson et al. 10.3389/fcosc.2023.1286459
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palaeoecological studies, which suggest that open, mosaic and fire-
adapted ecosystems are an integral part of many Malagasy
landscapes outside of the mesic northeast (Figure 1).
The “whole island forest”was an idea based on sparse data and
anecdotal evidence from the 18th century (McConnell and Kull,
2014). Nevertheless, deforestation rates are still measured relative to
a the presumed former 100% forest cover, leading to dramatic over-
estimation of deforestation rates in areas that harbor ancient open
and mosaic landscapes (Huff and Orengo, 2020). Such calculations
assume unforested landscapes are degraded, without investigation
the landscape history. This approach leads to neglect of the
conservation of open and mosaic landscapes and the ecosystem
services that they provide (Raveloaritiana et al., 2023) (see case
study 1). Such assumptions can also have negative consequences for
communities who are blamed for forest destruction or poor land
stewardship (Seagle, 2012) (see case study 2). It is therefore essential
to distinguish the extent of former forest cover prior to human
arrival in Madagascar, the historical accounts of forest cover and its
trajectory in recent centuries, in order that appropriate locations for
reforestation can be identified.
To accurately calculate deforestation rates, we need to
understand the former extent of forest cover in the early and
mid-Holocene. Prior to human settlement and expansion c 2500–
1350 years ago (Crowley, 2010;Douglass et al., 2019), closed canopy
rainforest occurred only in the mesic northeast of the island, and
eastern coastal strip (Straka, 1996;Moat and Smith, 2007). In the
north-west, where rainfall is lower and more seasonal, tropical dry
forests predominated, interspersed with succulents and heathlands
(shrublands dominated by the family Ericaceae) (Figure 2). In the
arid south-western part of the island, rainfall is too low to support
true forests (ie. those that are fire intolerant and with a closed or
near closed canopy) and open and arid-adapted landscapes
including the spiny thickets existed here (Figure 2). Between these
mesic and arid extremes, for example in the Highlands of Central
Madagascar, there were mosaic ecosystems where true forest
occurred only in patches (Figure 2) within matrices of ancient
fire-prone grasslands, heathlands and woodland or savannas (e.g.
(Burney et al., 2003;Virah-Sawmy et al., 2009b). These mosaic
landscapes need particular consideration in reforestation planning,
so that forests do not encroach on ancient open landscapes elements
that harbor unique biodiversity (Bond et al., 2008;Willis et al.,
2008). Specifically, fossil pollen data from lake sediments be used to
identify whether forest distribution was once continuous, or was
always patchy. This information can inform landscape management
and restoration plans, a point illustrated by a case study from the
Central Highlands of Madagascar.
2.1 Case study 1: Central Highlands
The Central Highlands are one of the main focuses for
afforestation schemes in Madagascar. They are currently
dominated by grasslands with patches of forest that are largely
confined to ravines. This landscape is assumed to be highly
degraded, however, palaeoecological investigations in the Central
Highlands suggest the long-term presence of mosaic landscapes,
where forest patches of variable extent co-existed with more open
landscape elements including grasslands, heathlands and savannas
FIGURE 1
Mosaic landscapes of Madagascar (A) next to Tsaramandroso village in the Northwest (Photo: A. Razafimanantsoa, 2011) (B) Landscape between
Morondava and Malaimbandy, Southwest (Photo: Andriantsaralaza, S. 2021) (C) Tampoketsa grassland –forest mosaic of the Central Highlands of
Madagascar (Photo: Kull, July 2019) (D) Littoral Forests, South East Efatsy Manombo, SE of Madagascar (Photo: Onjalalaina, 2020).
Gillson et al. 10.3389/fcosc.2023.1286459
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(Burney, 1987b;Gasse and Van Campo, 2001;Razafimanantsoa,
2022). Though rainfall in the Central Highlands is high enough to
support forests, open landscapes are likely maintained by fire and
herbivory, as occurs in much of sub-Saharan Africa (Bond et al.,
2005;Archibald et al., 2019). In such mosaic landscapes, co-
existence of different vegetation types in the same climatic zone is
mediated by topography, local hydrology and fire-vegetation-
herbivory feedbacks, with forest elements generally confined to
ravines which have greater water availability and are somewhat
protected from fire (Bond and Keeley, 2005;Bond et al., 2005;
Lehmann et al., 2011). Once established, fire sensitive forest
elements can maintain themselves by shading out the herbaceous
layer, which excludes fire except in extreme fire events or in the case
of deliberate clearance by people (Staver et al., 2011;Archibald et al.,
2019). Modeling experiments that predict forest cover based solely
on rainfall are therefore likely to over-estimate past and present
forest cover (Bond and Keeley, 2005;Bond et al., 2005) whereas
those including fire more accurately represent the occurrence of
savanna and grasslands throughout much of sub-Saharan Africa,
including all but the eastern-most coastal strip in Madagascar
(Bond and Keeley, 2005;Bond et al., 2005;Lehmann et al., 2011).
Fire- and grazing- adapted traits are common in many
highlands species, strongly suggesting the presence of open
vegetation elements, including grassland, heathland and savannas
(e.g. Burney, 1987a;Burney, 1987b;Straka, 1996;Razafimanantsoa,
2022). Some of the grasslands, especially on the western flank, are
highly biodiverse ancient grasslands, probably originating from the
Neogene period (Bond et al., 2008;Vorontsova et al., 2016;Hackel
et al., 2018;Vorontsova et al., 2020), while others are likely to be of
anthropogenic origin. A recent pollen record from Tampoketsa-
Ankazobe wetland, in the eastern slopes of the Highlands
demonstrated the abundance of ericoid shrubland during the
early Holocene, (Razafimanantsoa, 2022)(seeFigure 3)
confirming previous studies showing extensive presence of
heathlands (Burney, 1987c;Straka, 1996) and supported by the
high diversity of Ericaceae species in Madagascar (Grubb, 2003).
While attention usually focuses on forest loss, the conversion of
heathlands to grasslands warrants further attention and
conservation/restoration effort (Silander et al., 2023).
The palaeoecological and evolutionary data highlight the
importance of considering landscape-specific histories in
designing restoration in the Highlands. Though grasslands
dominate the Highlands today, it is likely that heathland,
woodland, savanna elements and forest patches were more
extensive in the past and co-existed alongside grasslands in some
areas (Burney, 1987a;Burney, 1987b;Straka, 1996;
Razafimanantsoa, 2022) (see Figure 3). A further concern of tree
planting initiatives is the selection of species, and how this selection
process takes place. Many forest plantation projects in the
Highlands use alien species that have negative effects on soil and
water and creating novel landscapes with reduced ecosystem service
provision. Eucalyptus has now infiltrated the remaining vestige of
FIGURE 2
Summary of the Holocene vegetation reconstruction in Madagascar based on published vegetation records. The time span covered by each record
is (A) Matsumoto and Burney, 1994;Burns et al., 2016;Voarintsoa et al., 2017;(B) Burney, 1993;Virah-Sawmy et al., 2016;Razanatsoa et al., 2021;
(C) Burney, 1987a;Burney, 1987b;Straka, 1996;Razafimanantsoa, 2022;Burney, 1987c;MacPhee et al., 1985;(D) Virah-Swamy et al., 2009a;2009b;
Virah-Sawmy, 2009. Map of main vegetation types based and simplified from Moat and Smith (2007).
Gillson et al. 10.3389/fcosc.2023.1286459
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indigenous Tapia savanna woodland (Uapaca bojeri), leading to
increased fire intensity and decreased regeneration of native species
including the Tapia seedlings (Baohanta et al., 2012;Rakotondrasoa
et al., 2012).
In assessing degradation and forest loss, the distinction is
critical between true forests from wooded savannas. The latter
maintain a near continuous herbaceous layer, with scattered tree
cover, and which are fire and grazing adapted. In the Central
Highlands, ancient Tapia (Uapaca) woodlands are now classified
as savannas based on their functional traits (Solofondranohatra
et al., 2018;Salmona et al., 2020). Native savanna trees such as the
“olive tree”Noronhia lowryi originated in the Miocene period, c. 6.5
million years ago, concurrent with the global spread of savannas
(Salmona et al., 2020). Classifying all savannas and heathlands as
degraded forests (see for example Joseph and Seymour, 2020;
Joseph and Seymour, 2021) is ecologically inaccurate and can lead
to further errors in estimating forest loss and in encouraging tree
planting schemes that replace unique open and mosaic ecosystems
and associated ecosystem services (Raveloaritiana et al., 2023).
Forest restoration (i.e. restoring previously forested areas with
indigenous species) should be emphasized over afforestation of
landscapes whose history is uncertain and which may contain
ancient grasslands, heathlands and savannas. In south-east of
Madagascar, for example, fast-growing native species such as
Treblus mauritianus,Syzygium bernieri,Treculia madagascariensis
and Uapaca thouarsii (Manjaribe et al., 2013) provide crucial
ecosystem services as well as maintaining biodiversity.
The false dichotomy between forest-grassland (Joseph and
Seymour, 2020;Joseph and Seymour, 2021) has diverted attention
away from the loss and conversion of heathlands and savannas in
the highlands, which are also ancient elements of the vegetation.
Distinguishing ancient grasslands and mosaic ecosystems from
derived anthropogenic ones is only one strand of a
comprehensive conservation approach to the highlands that
includes the conservation and restoration of ancient mosaic
landscapes, reforestation in appropriate areas, as well as fire
management that reflects the history and ecology of the region
(Razafimanantsoa, 2022).
3 Balancing trade-offs between
biodiversity, carbon storage a
resource management
The primary motivation for most forest plantation projects is
their perceived benefit to carbon storage to mitigate climate change.
For example, the global carbon mitigation frameworks of the Clean
Development Mechanism (CDM) and Reduced Emissions from
Deforestation and Degradation (REDD and REDD+) have created
new economic incentives for low income nations to engage in
afforestation in Africa (Leach and Scoones, 2015;Bastin et al., 2019).
However, in Madagascar, there is limited evidence for the success of
these projects (Neudert et al., 2017), attributed to the choice of
species, the way ecosystem service benefits are calculated, and to
whom, and how, benefits are accrued.
Carbon calculations are typically incomplete or inaccurate and
other ecosystem services, including cultural ones, are seldom
considered. A particular problem is tendency to use above ground
biomass as the main indicator of carbon stocks, which can lead to
the underestimation of carbon stored in open and mosaic
ecosystems. In Madagascar, as elsewhere, soil carbon pools hold
three times higher than above-ground (Powers et al., 2011;Grinand
et al., 2017;Ramifehiarivo et al., 2017;Dass et al., 2018;
Razafindrakoto et al., 2018;Veldman et al., 2019;Yang et al., 2019).
Payments for avoided deforestation can in theory provide a
more stable revenue than other forms of livelihood such as charcoal
production or swidden agriculture (Neudert et al., 2018). However,
communities are receiving fewer benefits than expected from REDD
projects due to structural and institutional challenges locally, and
also governance failure (Neudert et al., 2017). They therefore rely on
multiple coping strategies to buffer environmental shocks.
At the same time, there are competing interests for example
from agriculture or mining and these financial interests are backed
by powerful organisations that may overpower local voices and
concerns over biodiversity loss. While the benefits of carbon storage
and export revenue are derived globally and nationally, the costs of
biodiversity protection, including loss of agricultural areas, loss of
FIGURE 3
Summary pollen diagram of the dominant taxa in Tampoketsa-Ankazobe site, Central Highlands Madagascar during the Holocene (Adapted from
Razafimanantsoa, 2022).
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access to non-timber forest products, and loss of income derived
from charcoal production, are born locally (Scales, 2014;Neudert
et al., 2017).
For long-term sustainability, tangible compensations to local
communities based on principles of co-management and self-
determinacy are vital (Neudert et al., 2017;Bond et al., 2019;
Temperton et al., 2019;Razanatsoa et al., 2021). Collaborations
are needed that consider landscape history and social context in
restoration projects that improve food security and sustainability of
livelihoods, while also conserving the biodiversity on which
ecosystem services depend. Participatory and ground-up
approaches can help in selecting species that are locally
appropriate and therefore more likely to survive as well as
providing benefits beyond carbon storage, and material, cultural
and educational benefits that contribute to the sustainability of
conservation projects (Razanatsoa et al., 2021), especially where
competing interests are at play and valuable resources at stake
(Virah-Swamy et al., 2009a;2009b). Case study 2 from the Fort
Dauphin area of southeast Madagascar illustrates these points.
3.1 Case study 2: littoral forests of
southeast Madagascar
The littoral forests are Madagascar’s most endangered and range-
restricted vegetation type, and are of high conservation value due to
their scarcity and abundance of endemic species (Watson et al., 2004;
Ingram and Dawson, 2005;Consiglio et al., 2006;Dawson and
Ingram, 2008;Virah-Sawmy et al., 2009b). They are also used by
communities for a range ecosystem services including oil, firewood,
timber, medicine, animal food, human food, fibres and oil, as well as
for spiritual purposes (Dawson and Ingram, 2008;Virah-Sawmy
et al., 2014b). Their distribution closely coincides with the occurrence
of ilmenite, and the forest patches have therefore been targeted for
mining operations by QMM, a collaboration between Rio Tinto and
the Malagasy Government. In assessing the environmental impact of
mining operations, high rates of deforestation over recent decades
were calculated relative to a presumed 100% former forest cover (Huff
and Orengo, 2020). Based on this calculation, QMM argued that the
rate of littoral forest destruction by local communities was
unsustainable and would far exceed that of the planned mining
operations (Bidaud et al., 2015). This argument was key in the
progress and approval of the mining plans (Consiglio et al., 2006).
An alternative explanation for the patchy distribution of the
littoral forest is that its extent is controlled mainly by local
environmental factors such as soil and topography, and that it
already existed in a patchy or mosaic form before human
occupation. This hypothesis was tested by analyzing fossil pollen
from within and outside forest patches. The results showed the
presence of ancient fire adapted heathlands and open woodlands,
pre-dating human arrival in Madagascar (Virah-Sawmy et al.,
2009a;Virah-Sawmy et al., 2009b). Furthermore, the work
demonstrated the ability of the forests to recover from natural
disturbances including climatic perturbations and storm surges.
The results suggest that littoral forest was always fragmented, its
distribution confined to localized areas defined by local
topographic, edaphic and hydrological factors (Virah-Sawmy,
2009;Virah-Sawmy et al., 2009a;Virah-Sawmy et al., 2009b).
Forest loss has accelerated in recent decades, exacerbated by road
construction during the exploratory phases of mining operations
(Bidaud et al., 2015). However, the forest losses prior to mining
operations, labeled as ‘catastrophic’, were inferred from current
distribution and in incorrect assumption of 100% historical forest
cover. The falsely high deforestation rates were used to obscure the
negative impacts of mining, without conservation of the natural
mosaic composition of the landscape.
The findings have implications for how littoral forests are
managed and how the impacts of communities are perceived.
Though QMM planned for mining operations to have Net
Positive Impact (NPI), the success of this policy is disputed, and
the ilmenite operations have been criticized for their impact on
biodiversity and communities (Virah-Sawmy et al., 2014a;Huff and
Orengo, 2020). First, NPI is calculated relative to the projected rate
of forest destruction in the absence of mining, which as explained
above is erroneously high due to an assumed former 100% forest
cover and associated high rate of forest destruction. Any slowing of
this perceived deforestation rate (“avoided deforestation”)is
presented as a net positive impact i.e. forest destruction can still
occur, but if the rate is calculated to be lower than the current rate
calculated relative to past 100% forest cover, this is counted as a
positive impact. Second, the effect of other mining impacts included
construction of a dam, deforestation, afforestation with alien
species, destruction of culturally important sites, loss of access to
reeds and fisheries were not properly accounted for. Villagers
displaced from their land were poorly compensated, if at all, and
were moved to areas with soils unsuitable for farming and
dislocated from cultural heritage (Huff and Orengo, 2020).
Communities suffered loss of access to resources such reed beds
and fisheries, either directly through prohibition of access, or
indirectly through pollution of water and dam construction.
Education programs instigated by QMM have also been criticized,
doing little more than reinstate practices such as basket weaving
that were only disrupted because of mining operations (Huff, 2014;
Huff and Orengo, 2020). Rio Tinto retracted its NPI policy in 2016
on the grounds that it is not always achievable (de Silva et al., 2019).
Around Fort Dauphin, the meagre compensation, low land
price and loss of access to resources were accepted by poor
communities, who were struggling to survive. The high levels of
poverty and urgent need for employment and economic interests
and pressures by the national government and international
companies mean implementing strict conservation of all forests is
not an option (Virah-Swamy et al., 2009a;2009b). Instead, Virah-
Sawmy and Ebeling (2010) advocate for a more environmentally
sensitive and inclusive approach to conservation and landscape
management that considers biodiversity alongside local needs
cultural context and landscape history Virah-Sawmy and Ebeling
2009a;2009b. Palaecological information can help acknowledge or
hopefully reduce biases in the way we manage and restore
landscapes, especially when economic interests are at stake and
local communities can easily be marginalized. Nevertheless, the
spectre of “green-washing”remains ever-present in conservation
mechanisms that are embedded in neo-liberalism (Adams, 2017).
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4 Discussion –toward a
transdisciplinary approach
Though deforestation and forest fragmentation in Madagascar
are of great concern (Vieilledent et al., 2018), the misrepresentation
of ancient open or mosaic ecosystems as ‘degraded’forests or of
anthropogenic origin has hampered conservation protection,
alienated local communities and also enabled the destruction of
wildlife habitats due to mining endeavors and afforestation schemes
(Bond et al., 2008;Virah-Sawmy et al., 2014a;Bond et al., 2019;Huff
and Orengo, 2020). Such schemes have also contributed to the
disenfranchisement of communities and the loss of livelihoods
(Huff and Orengo, 2020). A transdisciplinary approach is needed
that considers landscape history, assesses ecosystem services
holistically and accurately, and includes stakeholders and
communities in decision-making and planning (Figure 4)(Fischer
et al., 2020;Edrisi and Abhilash, 2021;Robinson et al., 2021).
The perceptions, management and afforestation initiatives of
the Malagassy landscapes do not originate from local communities,
but are based on anecdotal “baselines”, reinforced by narratives
entangled in degradation and economic interests (Pollini, 2010;
Hajdu and Fischer, 2017). For example, the assumption that all
open ecosystems in Madagascar are degraded forests continues to
underpin calculations of deforestation rates and debates in the
literature (Huff and Orengo, 2020;Joseph and Seymour, 2020;
Joseph and Seymour, 2021). However, the dichotomy between
forest versus degraded forest is an artificial one that does not
consider the spectrum of open and mosaic landscapes including
savannas, heathlands and shrubland. Classifying such landscapes as
degraded forest leads to over-estimation of deforestation rates and
neglect of conservation of open and mosaic ecosystems including
ancient grasslands, heathlands and savanna woodlands (Bond
et al., 2019).
Current debates over whether open ecosystems are ancient or
anthropogenic in Madagascar (Joseph and Seymour, 2020;Joseph
and Seymour, 2021), are missing a key challenge for an informed
and bottom up approach to conservation and restoration, which is
to distinguish which open landscapes are ancient or anthropogenic,
their historic nature and their current value to communities in
ecosystem service provision (Vetter, 2020;Lehmann et al., 2021).
The current polarization of forested versus degraded land
obfuscates a sound ecological discussion of the ecological function
of open and mosaic landscapes and their longevity and importance
as biodiversity hotspots. Though principles for considering
landscape history in restoration ecology and reforestation are
emerging (Di Sacco et al., 2021), there have as yet been few
attempts to integrate long-term data sets into restoration plans. In
fact, a range of ecological and palaeoecological techniques are
available to explore these questions, yet they remain underutilized
(Virah-Sawmy et al., 2009b;Crowley et al., 2021;Gillson
et al., 2021).
Palaeoecology can provide important information on the
former extent of forest cover and the antiquity of mosaic and
open ecosystems. Palaeoecological data can also guide the species
used in reforestation, providing information on former forest
composition that could be used to select indigenous instead of
exotic species (Wingard et al., 2017). Though Pinus, Eucalyptus and
Acacia, are favored because of their high survival rate and rapid
growth compared with many native species (Carriere and
Randriambanona, 2007;Verhaegen et al., 2011;Kull et al., 2019;
Randriambanona et al., 2019), these plantations are low in
biodiversity and negatively impact soil and water provision
provided by open and mosaic ecosystems (Charles and Dukes,
2008;Stickler et al., 2009;Bond, 2019;Kull et al., 2019).
It is of course essential that the benefits of tree-planting on
carbon storage are accurately calculated. The benefits of
afforestation with non-native species are likely to be over-
estimated, when only above-ground biomass, the most commonly
used measurement in the REDD and REDD+ initiative, is used
(Leach and Scoones, 2015;Panfil and Harvey, 2016;Bastin et al.,
2019). Carbon calculations based on above-ground biomass have
underestimated carbon storage in grasslands, savannas and
heathlands, despite their potential as carbon sinks (Veldman
et al., 2019), favoring the accelerated loss of high-conservation-
value ecosystems with low above-ground biomass and exacerbating
loss of biodiversity elsewhere through displaced agricultural
expansion (Bond et al., 2019). In addition, it is critical that REDD
+ and other carbon storage and afforestation schemes consider the
wider range of ecosystem services provided by mosaic and open
ecosystems, and their importance in local livelihoods (Lehmann
et al., 2021). Some land-uses including agroforestry generally are
able to maintain good soil carbon stocks (Rakotovao et al., 2020).
Taviala (agroforestry) can help to maintain forest cover and also
provides food and other resources as well as opportunities for
producing high value cash crops such as vanilla and silk (Virah-
Sawmy et al., 2015;Reuter et al., 2016;Rakotonarivo et al., 2017;
FIGURE 4
An integrated assessment of landscapes that considers landscape
history, current ecological knowledge of pattern and process, and
stakeholder perspectives on the history, culture and ecosystem
services that landscapes provide.
Gillson et al. 10.3389/fcosc.2023.1286459
Frontiers in Conservation Science frontiersin.org07
Hending et al., 2018;Hewson et al., 2019;Hending et al., 2020;
Schüßler et al., 2020;Wurz et al., 2022). The benefits of agroforestry
to biodiversity are greater in forest-derived than fallow-derived land
(Osen et al., 2021) Multiple livelihood streams from Taviala can
buffer households against environmental shocks and improve the
security of land-tenure, as well as providing habitat for biodiversity
(Rakotonarivo et al., 2017;Schüßler et al., 2020).
Protected area expansion is another approach that can
potentially mitigate carbon emissions while at the same time
maintaining essential ecosystem services that bolster livelihoods,
especially during times of environmental and economic stress
(Waeber et al., 2016;Lehmann et al., 2021). However, it is
important that local communities are consulted in the designation
and management of such areas. In Madagascar, sacred groves
protected by local taboos were incorporated into protected area
planning (Virah-Sawmy et al., 2014b;Nopper et al., 2017;Schüßler
et al., 2020), so that conservation planning reinforced and did not
erode local culture and traditions.
Palaeoecology will be only one strand in developing a more
nuanced, ecologically sound and locally appropriate approach to
landscape restoration (Gillson et al., 2021). Local communities are
often the ones who benefit the least from afforestation, resource
extraction, and even some forms of conservation (Neudert
et al., 2017).
Understanding the importance of cultural landscapes in
maintaining diverse ecosystem services and buffering livelihoods
against environmental shocks will require close engagement with
communities. This will require a transdisciplinary approach that
incorporates more accurate calculation of carbon stocks, a better
understanding and reflexivity of landscape history, consideration of
process (how decisions are made) and impacts of access to resources
and livelihoods (Sayer et al., 2013;Fischer et al., 2020). A more
inclusive approach to restoration is needed that considers desired
futures and acceptable trade-offs between ecosystem services and
resource provision (Fischer et al., 2020;Di Sacco et al., 2021;Gillson
et al., 2021). Social processes are needed that aim to facilitate
constructive debate among interest groups toward a common
understanding and resolution of complex objectives in
restoration, especially when there are competing powerful
economic interests at stake (Sayer et al., 2013;Fischer et al.,
2020). Palaeoecological information has the potential to be an
important source of data that can help acknowledge or reduce
biases in the way we perceive, manage and restore landscapes
(Gillson, 2015).
5 Conclusions
Though motivated by legitimate concerns to halt deforestation
and mitigate global emissions, poorly planned and implemented tree-
planting projects risk - and have already resulted in - exacerbated
degradation either directly through loss or replacement of indigenous
vegetation or indirectly through social and economic displacement
(Fairhead et al., 2012;Larson et al., 2013).
In Madagascar, accumulated evolutionary, comparative and
palaeoecological knowledge suggest the antiquity of open and
mosaic ecosystems, establishing them as valid conservation targets
(Burney et al., 2003;Bond et al., 2008;Virah-Sawmy et al., 2009b;
Vorontsova et al., 2016;Hackel et al., 2018;Vorontsova et al., 2020).
Mosaic landscapes of forested and open ecosystems provide higher
biodiversity and a wider range of ecosystem services than forested
landscapes alone (Raveloaritiana et al., 2023)–and this is especially
true where afforestation schemes use invasive alien species. The
carbon storage capacity of open and mosaic landscapes has been
underestimated and needs to be considered in restoration projects.
Looking ahead, it is vital for both biodiversity and local
communities that carbon storage schemes look beyond
afforestation to consider more complex, nuanced and authentic
landscape mosaics that reflect landscape history and better serve
both biodiversity and people. Understanding the history and culture
of landscapes requires a multi-disciplinary approach underpinned
by biological science but that considers local context, landscape
history and stakeholder needs to co-design restoration,
conservation and carbon mitigation projects that contribute to
social-ecological resilience.
Author contributions
LG: Conceptualization, Funding acquisition, Writing –original
draft, Writing –review & editing. ER: Conceptualization,
Visualization, Writing –review & editing. AR: Conceptualization,
Data curation, Formal Analysis, Visualization, Writing –review &
editing. MV: Writing –review & editing. AE: Writing –review
& editing.
Funding
The author(s) declare financial support was received for the
research, authorship, and/or publication of this article. The authors
gratefully acknowledge the following sources of funding which
contributed to this work: National Research Foundation (NRF)
Competitive Programme for Rated Researchers (Grant Number
118538), NRF/African Origins Platform (Grant Number 117666),
NRF/Global Change Grand Challenge/SASSCAL (Grant number
118589), and the University of Cape Town Vice Chancellor’s Future
Leaders Fund.
Conflict of interest
The authors declare that the research was conducted in the
absence of any commercial or financial relationships that could be
construed as a potential conflict of interest.
The author(s) declared that they were an editorial board
member of Frontiers, at the time of submission. This had no
impact on the peer review process and the final decision.
Gillson et al. 10.3389/fcosc.2023.1286459
Frontiers in Conservation Science frontiersin.org08
Publisher’s note
All claims expressed in this article are solely those of the authors
and do not necessarily represent those of their affiliated
organizations, or those of the publisher, the editors and the
reviewers. Any product that may be evaluated in this article, or
claim that may be made by its manufacturer, is not guaranteed or
endorsed by the publisher.
References
Adams, W. (2017). Sleeping with the enemy? Biodiversity conservation, corporations
and the green economy. J. Pol. Ecol. 24 (1), 243–257. doi: 10.2458/v24i1.20804
Antonelli, A., Smith, R. J., Perrigo, A. L., Crottini, A., Hackel, J., Testo, W., et al.
(2022). Madagascar’s extraordinary biodiversity: Evolution, distribution, and use.
Science 378, eabf0869. doi: 10.1126/science.abf0869
Archibald, S., Bond, W. J., Hoffmann, W., Lehmann, C., Staver, C., and Stevens, N.
(2019). “Distribution and determinants of savannas,”in Savanna woody plants and
large herbivores. Eds. P. F. Scogings and M. Sankaran (Chichester: Wiley), 1–24.
Baohanta, R., Thioulouse, J., Ramanankierana, H., Prin, Y., Rasolomampianina, R.,
Baudoin, E., et al. (2012). Restoring native forest ecosystems after exotic tree plantation
in Madagascar: combination of the local ectotrophic species Leptolena bojeriana and
Uapaca bojeri mitigates the negative influence of the exotic species Eucalyptus
camaldulensis and Pinus patula. Biol. Invasions 14, 2407–2421. doi: 10.1007/s10530-
012-0238-5
Barak, R. S., Hipp, A. L., Cavender-Bares, J., Pearse, W. D., Hotchkiss, S. C., Lynch, E.
A., et al. (2016). Taking the long view: integrating recorded, archeological,
paleoecological, and evolutionary data into ecological restoration. Int. J. Plant Sci.
177, 90–102. doi: 10.1086/683394
Bastin, J.-F., Finegold, Y., Garcia, C., Mollicone, D., Rezende, M., Routh, D., et al.
(2019). The global tree restoration potential. Science 365, 76–79. doi: 10.1126/
science.aax0848
Baumann, F. (2021). The next frontier—Human development and the anthropocene:
UNDP human development report 2020. Environment: Sci. Policy Sustain. Dev. 63, 34–
40. doi: 10.1080/00139157.2021.1898908
Bidaud, C.,Hrabanski, M., and Meral, P. (2015). Voluntary biodiversity offset strategies
in Madagascar. Ecosystem Serv. 15, 181–189. doi: 10.1016/j.ecoser.2015.02.011
Boissière, M., Atmadja, S., Guariguata, M. R., Kassa, H., and Sist, P. (2021). Perspectives
on the socio-economic challenges and opportunities for tree planting: A case study of
Ethiopia. For. Ecol. Manage. 497, 119488. doi: 10.1016/j.foreco.2021.119488
Bond, W. J. (2019). Open ecosystems: ecology and evolution beyond the forest edge
(Oxford: Oxford University Press).
Bond, W. J., and Keeley, J. E. (2005). Fire as a global 'herbivore': the ecology and
evolution of flammable ecosystems. Trends Ecol. Evol. 20, 387–394. doi: 10.1016/
j.tree.2005.04.025
Bond, W. J., Silander, J. A. Jr., Ranaivonasy, J., and Ratsirarson, J. (2008). The
Antiquity of Madagascar's grasslands and the rise of C4 biomes. J. Biogeography 35,
1743–1758. doi: 10.1111/j.1365-2699.2008.01923.x
Bond, W. J., Stevens, N., Midgley, G. F., and Lehmann, C. E. (2019). The trouble with
trees: afforestation plans for Africa. Trends Ecol. Evol. 34, 963–965. doi: 10.1016/
j.tree.2019.08.003
Bond, W. J., Woodward, F. I., and Midgley, G. F. (2005). The global distribution of
ecosystems in a world without fire. New Phytol. 165, 525–538. doi: 10.1111/j.1469-
8137.2004.01252.x
Brancalion, P. H., and Holl, K. D. (2020). Guidance for successful tree planting
initiatives. J. Appl. Ecol. 57, 2349–2361. doi: 10.1111/1365-2664.13725
Bredemeier, M., and Dohrenbusch, A. (2009). Afforestation and reforestation.
Biodiversity: Structure Function-Volume II 2, 219.
Bulte, E. H., Lipper, L., Stringer, R., and Zilberman, D. (2008). Payments for
ecosystem services and poverty reduction: concepts, issues, and empirical
perspectives. Environ. Dev. Economics 13, 245–254. doi: 10.1017/S1355770X08004348
Burney, D. A. (1987a). “Pre-settlement vegetation changes at Lake Tritrivakely,
Madagascar,”Palaeoecology of Africa and the surrounding islands 18, 357–381.
Burney, D. A. (1987b). Late Holocene vegetational change in central Madagascar.
Quaternary Res. 28, 130–143. doi: 10.1016/0033-5894(87)90038-X
Burney, D. (1987c). Late Quaternary Stratigraphic Charcoals Records from
Madagascar. Quaternary Res. 28:274–280.
Burney, D. A. (1993). Recent animal extinctions; recipes for disaster. Am. Scientist 81,
530–541.
Burney, D. A., Robinson, G. S., and Burney, L. P. (2003). Sporormiella and the late
Holocene extinctions in Madagascar. Proc. Natl. Acad. Sci. 100, 10800–10805.
doi: 10.1073/pnas.1534700100
Burns, S. J., Godfrey, L. R., Faina, P., McGee, D., Hardt, B., Ranivoharimanana, L.,
et al. (2016). Rapid human-induced landscape transformation in madagascar at the end
of the first millennium of the common era. Quaternary Sci. Rev. 134, 92–99.
Carriere, S. M., and Randriambanona, H. (2007). Biodiversiteintroduite et
autochtone: antagonisme ou complementarite? Le cas de l'eucalyptus à Madagascar.
BOIS FORETS Des. TROPIQUES 292, 5–21. doi: 10.19182/bft2007.292.a20349
Charles, H., and Dukes, J. S. (2008). “Impacts of invasive species on ecosystem
services,”in Biological invasions (Berlin, Heidelberg: Springer), 217–237.
Consiglio, T., Schatz, G. E., McPherson, G., Lowry, P. P., Rabenantoandro, J., Rogers,
Z. S., et al. (2006). Deforestation and plant diversity of Madagascar's littoral forests.
Conserv. Biol. 20, 1799–1803. doi: 10.1111/j.1523-1739.2006.00562.x
Crowley, B. E. (2010). A refined chronology of prehistoric Madagascar and the
demise of the megafauna. Quaternary Sci. Rev. 29, 2591–2603. doi: 10.1016/
j.quascirev.2010.06.030
Crowley, B. E., Godfrey, L. R., Hansford, J. P., and Samonds, K. E. (2021). Seeing the
forest for the trees—and the grasses: revisiting the evidence for grazer-maintained
grasslands in Madagascar's Central Highlands. Proc.R.Soc.B288, 20201785.
doi: 10.1098/rspb.2020.1785
Dass, P., Houlton, B. Z., Wang, Y., and Warlind, D. (2018). Grasslands may be more
reliable carbon sinks than forests in California. Environ. Res. Lett. 13, 074027.
doi: 10.1088/1748-9326/aacb39
Dawson, T. P., and Ingram, J. C. (2008). Sustainable livelihoods and forest resources
in Madagascar: A multi-scale analysis using remote sensing. Environ. Sci. 5, 129–143.
doi: 10.1080/15693430801912188
de Silva, G. C., Regan, E. C., Pollard, E. H. B., and Addison, P. F. E. (2019). The
evolution of corporate no net loss and net positive impact biodiversity commitments:
Understanding appetite and addressing challenges. Business Strategy Environ. 28,
1481–1495. doi: 10.1002/bse.2379
Dietl, G. P., Kidwell, S. M., Brenner, M., Burney, D. A., Flessa, K. W., Jackson, S. T.,
et al. (2015). Conservation paleobiology: leveraging knowledge of the past to inform
conservation and restoration. Annu. Rev. Earth Planetary Sci. 43, 79–103. doi: 10.1146/
annurev-earth-040610-133349
Di Sacco, A., Hardwick, K. A., Blakesley, D., Brancalion, P. H., Breman, E., Cecilio
Rebola, L., et al. (2021). Ten golden rules for reforestation to optimize carbon
sequestration, biodiversity recovery and livelihood benefits. Global Change Biol. 27,
1328–1348. doi: 10.1111/gcb.15498
Douglass, K., Hixon, S., Wright, H. T., Godfrey, L. R., Crowley, B. E., Manjakahery,
B., et al. (2019). A critical review of radiocarbon dates clarifies the human settlement of
Madagascar. Quaternary Sci. Rev. 221, 105878. doi: 10.1016/j.quascirev.2019.105878
Edrisi, S. A., and Abhilash, P. (2021). Need of transdisciplinary research for
accelerating land restoration during UN decade on ecosystem restoration. Restor.
Ecol. 29 (8), e13531. doi: 10.1111/rec.13531
Fairhead, J., Leach, M., and Scoones, I. (2012). Green Grabbing: a new appropriation
of nature? J. Peasant Stud. 39, 237–261. doi: 10.1080/03066150.2012.671770
Falk, D. A., Watts, A. C., and Thode, A. E. (2019). Scaling ecological resilience. Front.
Ecol. Evol. 7. doi: 10.3389/fevo.2019.00275
Fischer, J., Riechers, M., Loos, J., Martin-Lopez, B., and Temperton, V. M. (2020).
Making the UN Decade on Ecosystem Restoration a social-ecological endeavour.
Trends Ecol. Evol. 36 (1), 20–28. doi: 10.1016/j.tree.2020.08.018
Garcı
a-Dory, F., Houzer, E., and Scoones, I. (2021). Livestock and climate justice:
Challenging mainstream policy narratives. IDS Bulletin 53 (4), 47–64. doi: 10.19088/
1968-2021.128
Gasse, F., and Van Campo, E. (2001). Late Quaternary environmental changes from a
pollen and diatom record in the soutehrn tropics (Lake Tritivakely, Madagascar).
Palaeogeography Palaeoclimatology Palaeoecol. 167, 287–308. doi: 10.1016/S0031-0182
(00)00242-X
Gillson, L. (2015). Biodiversity Conservation and Environmental Change: Using
palaeoecology to manage dynamic landscapes in the Anthropocene (Oxford, U.K:
Oxford University Press).
Gillson, L., Dirk, C., and Gell, P. (2021). Using long-term data to inform a decision
pathway for restoration of ecosystem resilience. Anthropocene 36, 100315. doi: 10.1016/
j.ancene.2021.100315
Grinand, C., Le Maire, G., Vieilledent, G., Razakamanarivo, H., Razafimbelo, T., and
Bernoux, M. (2017). Estimating temporal changes in soil carbon stocks at ecoregional
scale in Madagascar using remote-sensing. Int. J. Appl. Earth observation
geoinformation 54, 1–14. doi: 10.1016/j.jag.2016.09.002
Gross-Camp, N. D., Martin, A., McGuire, S., Kebede, B., and Munyarukaza, J. (2012).
Payments for ecosystem services in an African protected area: exploring issues of
Gillson et al. 10.3389/fcosc.2023.1286459
Frontiers in Conservation Science frontiersin.org09
legitimacy, fairness, equity and effectiveness. Oryx 46, 24–33. doi: 10.1017/
S0030605311001372
Hackel, J., Vorontsova, M. S., Nanjarisoa, O. P., Hall, R. C., Razanatsoa, J., Malakasi,
P., et al. (2018). Grass diversification in Madagascar: in situ radiation of two large C3
shade clades and support for a Miocene to Pliocene origin of C4 grassy biomes. J.
Biogeography 45, 750–761. doi: 10.1111/jbi.13147
Hajdu, F., and Fischer, K. (2017). Problems, causes and solutions in the forest carbon
discourse: a framework for analysing degradation narratives. Climate Dev. 9, 537–547.
doi: 10.1080/17565529.2016.1174663
Hending, D., Andrianiaina, A., Maxfield, P., Rakotomalala, Z., and Cotton, S. (2020).
Floral species richness, structural diversity and conservation value of vanilla
agroecosystems in Madagascar. Afr. J. Ecol. 58, 100–111. doi: 10.1111/aje.12682
Hending, D., Andrianiaina, A., Rakotomalala, Z., and Cotton, S. (2018). The use of
vanilla plantations by lemurs: encouraging findings for both lemur conservation and
sustainable agroforestry in the Sava region, northeast Madagascar. Int. J. Primatology
39, 141–153. doi: 10.1007/s10764-018-0022-1
Hewson, J., Razafimanahaka, J. H., Wright, T. M., Mandimbiniaina, R., Mulligan, M.,
Jones, J. P., et al. (2019). Land change modelling to inform strategic decisions on Forest
cover and CO 2 emissions in eastern Madagascar. Environ. Conserv. 46, 25–33.
doi: 10.1017/S0376892918000358
Huff, A. (2014). Weathering the'Long Wounded Year': livelihoods, nutrition and
changing political ecologies in the Mikea Forest Region, Madagascar. J. Political Ecol.
21, 83–107.
Huff, A., and Orengo, Y. (2020). Resource warfare, pacification and the spectacle of
‘green’development: Logics of violence in engineering extraction in southern
Madagascar. Political Geogr. 81, 102195. doi: 10.1016/j.polgeo.2020.102195
Ingram, J. C., and Dawson, T. P. (2005). Climate change impacts and vegetation
response on the island of Madagascar. Philos. Trans. R. Soc. A: Mathematical Phys. Eng.
Sci. 363, 55–59. doi: 10.1098/rsta.2004.1476
Joseph, G. S., and Seymour, C. L. (2020). Madagascan highlands: originally woodland
and forest containing endemic grasses, not grazing-adapted grassland. Proc. R. Soc. B
287, 20201956. doi: 10.1098/rspb.2020.1956
Joseph, G. S., and Seymour, C. L. (2021). The unlikely’Antiquity of Madagascar's
grasslands’: Disproportionately forest-limited endemic fauna support anthropogenic
transformation from woodland. J. Biogeogr. 48 (8), 2111–2115. doi: 10.1111/jbi.14132
Kronenberg, J., and Hubacek, K. (2013). Could payments for ecosystem services
create an “ecosystem service curse”?Ecol. Soc. 18 (1), 10. doi: 10.5751/ES-05240-180110
Kull, C. A., Harimanana, S. L., Andrianoro, A. R., and Rajoelison, L. G. (2019).
Divergent perceptions of the ‘neo-Australian’forests of lowland eastern Madagascar:
Invasions, transitions, and livelihoods. J. Environ. Manage. 229, 48–56. doi: 10.1016/
j.jenvman.2018.06.004
Lacroix, E., Carodenuto, S., Richter, F., Pistorius, T., and Tennigkeit, T. (2016).
Restauration des paysages forestiers Evaluation des potentialites dans le contexte des
engagements de Bonn 2.0 et de la Declaration de New York sur les forets. Methodologie
resultats pour Madagascar. Mars. Available at: https://afr100.org/sites/default/files/
2022-11/06-MDG_RPF_MEOR_Madagascar_Final.pdf.
Larson, A. M., Brockhaus, M., Sunderlin, W. D., Duchelle, A., Babon, A., Dokken, T.,
et al. (2013). Land tenure and REDD+: The good, the bad and the ugly. Global Environ.
Change 23, 678–689. doi: 10.1016/j.gloenvcha.2013.02.014
Leach, M., and Scoones, I. (2015). “Political ecologies of carbon in Africa,”in Carbon
conflicts and forest landscapes in Africa, M. Leach and I. Scoones (eds) (London:
Routledge), 21–62.
Lehmann, C. E. R., Archibald, S. A., Hoffmann, W. A., and Bond, W. J. (2011).
Deciphering the distribution of the savanna biome. New Phytol. 191, 197–209.
doi: 10.1111/j.1469-8137.2011.03689.x
Lehmann, C. E., Solofondranohatra, C. L., and Vorontsova, M. S. (2021). Beyond
ancient versus anthropogenic for Madagascar's grassy ecosystems. A Reply to: Crowley
et al.(2021). Proc. R. Soc. B 288, 20210388. doi: 10.1098/rspb.2021.0388
MacPhee, R., Burney, D., and Wells, N. (1985). Early holocene chronology and
environment of ampasambazimba, a malagasy subfossil lemur site. Int. J. Primatology 6,
463–489.
Mandimbisoa, R. (2021). Une vitrine presidentielle de reboisement à Analamanga.
(Antananarivo: https://Madagascar-Tribune.com)
Manjaribe, C., Frasier, C. L., Rakouth, B., and Louis, E. E. (2013). Ecological
restoration and reforestation of fragmented forests in Kianjavato, Madagascar. Int. J.
Ecol, 726275. doi: 10.1155/2013/726275
Matsumoto, K., and Burney, D. A. (1994). Late holocene environments at lake
mitsinjo, northwestern madagascar. Holocene 4, 16–24.
McConnell, W. J., and Kull, C. A. (2014). “Deforestation in Madagascar: Debates
over the island's forest cover and challenges of measuring forest change,”in
Conservation and environmental management in Madagascar, Ed, I. R. Scales (New
York and London: Routledge), 67–104.
Moat, J., and Smith, P. (2007). Atlas of the Vegetation of Madagascar (London, UK:
The Royal Botanic Gardens, Kew. Kew Publishing).
Myers, N., Mittermeier, R. A., Mittermeier, C. G., da Fonseca, G. A. B., and Kents, J.
(2000). Biodiversity hotspots for conservation priorities. Nature 403, 853–858.
doi: 10.1038/35002501
Neudert, R., Ganzhorn, J. U., and WÄTzold, F. (2017). Global benefits and local costs
–The dilemma of tropical forest conservation: A review of the situation in Madagascar.
Environ. Conserv. 44, 82–96. doi: 10.1017/S0376892916000552
Neudert, R., Olschofsky, K., Kübler, D., Prill, L., Köhl, M., and Wätzold, F. (2018).
Opportunity costs of conserving a dry tropical forest under REDD+: The case of the
spiny dry forest in southwestern Madagascar. For. Policy Economics 95, 102–114.
doi: 10.1016/j.forpol.2018.07.013
Neugarten, R. A., Honzak, M., Carret, P., Koenig, K., Andriamaro, L., Cano, C. A.,
et al. (2016). Rapid assessment of ecosystem service co-benefits of biodiversity priority
areas in Madagascar. PLoS One 11, e0168575. doi: 10.1371/journal.pone.0168575
Nopper, J., Ranaivojaona, A., Riemann, J. C., Rödel, M.-O., and Ganzhorn, J. U.
(2017). One forest is not like another: The contribution of community-based natural
resource management to reptile conservation in Madagascar. Trop. Conserv. Sci. 10,
1940082917693234. doi: 10.1177/1940082917693234
Osen, K., Soazafy, M. R., Martin, D. A., Wurz, A., März, A., Ranarijaona, H. L. T.,
et al. (2021). Land-use history determines stand structure and tree diversity in vanilla
agroforests of northeastern Madagascar. Appl. Vegetation Sci. 24, e12563. doi: 10.1111/
avsc.12563
Panfil, S. N., and Harvey, C. A. (2016). REDD+ and biodiversity conservation: A
review of the biodiversity goals, monitoring methods, and impacts of 80 REDD+
projects. Conserv. Lett. 9, 143–150. doi: 10.1111/conl.12188
Pollini, J. (2010). Environmental degradation narratives in Madagascar: From
colonial hegemonies to humanist revisionism. Geoforum 41, 711–722. doi: 10.1016/
j.geoforum.2010.04.001
Powers, J. S., Corre, M. D., Twine, T. E., and Veldkamp, E. (2011). Geographic bias of
field observations of soil carbon stocks with tropical land-use changes precludes spatial
extrapolation. Proc. Natl. Acad. Sci. 108, 6318–6322. doi: 10.1073/pnas.1016774108
Rakotonarivo, O. S., Jacobsen, J. B., Larsen, H. O., Jones, J. P., Nielsen, M. R.,
Ramamonjisoa, B. S., et al. (2017). Qualitative and quantitative evidence on the true
local welfare costs of forest conservation in Madagascar: are discrete choice experiments
a valid ex ante tool? World Dev. 94, 478–491. doi: 10.1016/j.worlddev.2017.02.009
Rakotondrasoa, O. L., Malaisse, F., Rajoelison, G. L., Razafimanantsoa, T. M.,
Rabearisoa, M. R., Ramamonjisoa, B. S., et al. (2012). Tapia forest, endemic
ecosystem to Madagascar: ecology, functions, causes of degradation and
transformation: a review. Biotechnologie Agronomie SocieteEnvironnement 16, 541–
552.
Rakotovao, N. H., Ramifehiarivo, N., Grinand, C., Razakamanarivo, H. R., Albrecht,
A., and Razafimbelo, T. M. (2020). Impact of agroecological practices on soil organic
carbon sequestration using synchronic and diachronic approaches in Madagascar. J.
Agric. Rural Dev. Tropics Subtropics (JARTS) 121, 113–125.
Ramifehiarivo, N., Brossard, M., Grinand, C., Andriamananjara, A., Razafimbelo, T.,
Rasolohery, A., et al. (2017). Mapping soil organic carbon on a national scale: Towards
an improved and updated map of Madagascar. Geoderma Regional 9, 29–38.
doi: 10.1016/j.geodrs.2016.12.002
Randriambanona, H., Randriamalala, J. R., and Carrière, S. M. (2019). Native forest
regeneration and vegetation dynamics in non-native Pinus patula tree plantations in
Madagascar. For. Ecol. Manage. 446, 20–28. doi: 10.1016/j.foreco.2019.05.019
Ranjatson, P., Rebecca, M., Randrianasolo, R., Mananga, J., Razafimbelo, N., and
Steven, L. (2019). “Tenure security and forest landscape restoration: Results from
exploratory research in Boeny, Madagascar,”in World bank conference on land and
poverty (Washington DC: The World Bank), 1–28. Available at: https://events.
globallandscapesforum.org/wp-content/uploads/sites/2/2019/10/PMcLain1901.pdf
Raveloaritiana, E., Wurz, A., Osen, K., Soazafy, M. R., Grass, I., Martin, D. A., et al.
(2023). Complementary ecosystem services from multiple land uses highlight the
importance of tropical mosaic landscapes. Ambio 52, 1558–1574. doi: 10.1007/s13280-
023-01888-3
Razafimanantsoa, A. H. I. (2022). Holocene vegetation history and environmental
change in the forest-grassland mosaic of the Central Highlands of Madagascar (Cape
Town: Doctoral. University of Cape Town).
Razafindrakoto, M., Andriamananjara, A., Razafimbelo, T., Hewson, J., Andrisoa, R.
H., Jones, J. P. G., et al. (2018). “Chapter 3 - organic carbon stocks in all pools following
land cover change in the rainforest of Madagascar,”in Soil Management and Climate
Change. Eds. MA
. Muñoz and R. Zornoza (Cambridge, Massachusetts, United States:
Academic Press), 25–37.
Razanatsoa,E.,Andriantsaralaza,S.,Holmes,S.M.,Rakotonarivo,O.S.,
Ratsifandrihamanana, A. N., Randriamiharisoa, L., et al. (2021). Fostering local
involvement for biodiversity conservation in tropical regions: Lessons from
Madagascar during the COVID-19 pandemic. Biotropica 53 (4), 994–1003. doi:
10.1111/btp.12967
Reuter, K. E., Randell, H., Wills, A. R., and Sewall, B. J. (2016). The consumption of
wild meat in Madagascar: drivers, popularity and food security. Environ. Conserv. 43,
273. doi: 10.1017/S0376892916000059
Rid, T. (2010). The nineteenth century origins of counterinsurgency doctrine. J.
Strategic Stud. 33, 727–758. doi: 10.1080/01402390.2010.498259
Robinson, J. M., Gellie, N., MacCarthy, D., Mills, J. G., O'Donnell, K., and Redvers, N.
(2021). Traditional ecological knowledge in restoration ecology: a Call to listen deeply,
to engage with, and respect indigenous voices. Restor. Ecol. 29, e13381. doi: 10.1111/
rec.13381
Gillson et al. 10.3389/fcosc.2023.1286459
Frontiers in Conservation Science frontiersin.org10
Salmona,J.,Olofsson,J.K.,Hong-Wa,C.,Razanatsoa,J.,Rakotonasolo,F.,
Ralimanana, H., et al. (2020). Late Miocene origin and recent population collapse of
the Malagasy savanna olive tree (Noronhia lowryi). Biol. J. Linn. Soc. 129, 227–243.
doi: 10.1093/biolinnean/blz164
Sayer, J., Sunderland, T., Ghazoul, J., Pfund, J.-L., Sheil, D., Meijaard, E., et al. (2013).
Ten principles for a landscape approach to reconciling agriculture, conservation, and
other competing land uses. Proc. Natl. Acad. Sci. 110, 8349–8356. doi: 10.1073/
pnas.1210595110
Scales, I. R. (2014). The future of conservation and development in Madagascar: time
for a new paradigm? Madagascar Conserv. Dev. 9, 5–12. doi: 10.4314/mcd.v9i1.2
Schüßler, D., Mantilla-Contreras, J., Stadtmann, R., Ratsimbazafy, J. H., and
Radespiel, U. (2020). Identification of crucial stepping stone habitats for biodiversity
conservation in northeastern Madagascar using remote sensing and comparative
predictive modeling. Biodiversity Conserv. 29, 2161–2184. doi: 10.1007/s10531-020-
01965-z
Seagle, C. (2012). Inverting the impacts: Mining, conservation and sustainability
claims near the Rio Tinto/QMM ilmenite mine in Southeast Madagascar. J. Peasant
Stud. 39, 447–477. doi: 10.1080/03066150.2012.671769
Silander, J. A. Jr., Bond, W. J., and Ratsirarson, J. (2023). The grassy ecosystems of
Madagascar in context: Ecology, evolution, and conservation. Plants People Planet.
doi: 10.1002/ppp3.10417
Solofondranohatra, C. L., Vorontsova, M. S., Hackel, J., Besnard, G., Cable, S.,
Williams, J., et al. (2018). Grass functional traits differentiate forest and savanna in the
Madagascar central highlands. Front. Ecol. Evol. 6, 184. doi: 10.3389/fevo.2018.00184
Staver,A.C.,Archibald,S.,andLevin,S.A.(2011).Theglobalextentand
determinants of Savanna and forest as alternative biome states. Science 334, 230–232.
doi: 10.1126/science.1210465
Stickler, C. M., Nepstad, D. C., Coe, M. T., McGrath, D. G., Rodrigues, H. O., Walker,
W. S., et al. (2009). The potential ecological costs and cobenefits of REDD: a critical
review and case study from the Amazon region. Global Change Biol. 15, 2803–2824.
doi: 10.1111/j.1365-2486.2009.02109.x
Straka, H. (1996). “Histoire de la vegetation de Madagascar oriental dans les derniers
100 millenaires,”in BiogeHographie de Madagascar (Paris: ORSTOM), 37.
Suzzi-Simmons, A. (2023). Status of deforestation of Madagascar. Global Ecol.
Conserv. 42, e02389. doi: 10.1016/j.gecco.2023.e02389
Temperton, V. M., Buchmann, N., Buisson, E., Durigan, G., Kazmierczak, Ł., Perring,
M. P., et al. (2019). Step back from the forest and step up to the Bonn Challenge: how a
broad ecological perspective can promote successful landscape restoration. Restor. Ecol.
27, 705–719. doi: 10.1111/rec.12989
Tsayem Demaze, M. (2014). L’enrolement de Madagascar dans la REDD+:
domestiquer une opportuniteinternationale. VertigO: la revue electronique en sciences
de l’environnement. (Open Edition Journals) 14 (1).
Veldman, J. W., Aleman, J. C., Alvarado, S. T., Anderson, T. M., Archibald, S., Bond,
W. J., et al. (2019). Comment on “The global tree restoration potential”.Science 366,
eaay7976. doi: 10.1126/science.aay7976
Veldman, J. W., Overbeck, G., Negreiros, D., Mahy, G., Le Stradic, S., Fernandes, G.
W., et al. (2015). Tyranny of trees in grassy biomes. Science 347, 484–485. doi: 10.1126/
science.347.6221.484-c
Verhaegen, D., Randrianjafy, H., Montagne, P., Danthu, P., Rabevohitra, R., Tassin,
J., et al. (2011). Reconstructing the history of Eucalyptus genus introduction in
Madagascar. Bois Forets Des. Tropiques 309 (3), 18–25. doi: 10.19182/
bft2011.309.a20462
Vetter, S. (2020). With power comes responsibility–A rangelands perspective on
forest landscape restoration. Front. Sustain. Food Syst. 4, 225. doi: 10.3389/
fsufs.2020.549483
Vieilledent, G., Grinand, C., Rakotomalala, F. A., Ranaivosoa, R., Rakotoarijaona, J.-
R., Allnutt, T. F., et al. (2018). Combining global tree cover loss data with historical
national forest cover maps to look at six decades of deforestation and forest
fragmentation in Madagascar. Biol. Conserv. 222, 189–197. doi: 10.1016/
j.biocon.2018.04.008
Virah-Sawmy, M. (2009). Ecosystem management in Madagascar during global
change. Conserv. Lett. 2, 163–170. doi: 10.1111/j.1755-263X.2009.00066.x
Virah-Sawmy, M., and Ebeling, J. (2010). The difficult road toward real-world
engagement: conservation science and mining in southern Madagascar. Conserv.
Lett. 3, 288–289. doi: 10.1111/j.1755-263X.2010.00126.x
Virah-Sawmy, M., Ebeling, J., and Taplin, R. (2014a). Mining and biodiversity offsets:
A transparent and science-based approach to measure “no-net-loss”.J. Environ.
Manage. 143, 61–70. doi: 10.1016/j.jenvman.2014.03.027
Virah-Sawmy, M., Gardner, C., and Ratsifandrihamanana, A. (2014b). “The Durban
vision in practice. Experiences in the participatory governance of Madagascar's new
protected areas,”in Conservation and Environmental Management in Madagascar. Ed.
I. Scales (New York and London: Routledge), 216–251.
Virah-Sawmy, M., Gillson, L., Gardner, C., Anderson, A., Clark, G., and Haberle, S.
(2015). A landscape vulnerability framework for identifying integrated conservation
and adaptation pathways to climate change: the case of Madagascar’s spiny forest.
Landscape Ecol. 31, 637–654. doi: 10.1007/s10980-015-0269-2
Virah-Sawmy, M., Gillson, L., and Willis, K. J. (2009a). How does spatial
heterogeneity influence resilience to climatic changes? Ecol. dynamics southeast
Madagascar. Ecol. Monogr. 79, 557–574. doi: 10.1890/08-1210.1
Virah-Sawmy, M., Willis, K. J., and Gillson, L. (2009b). Threshold response of
Madagascar's littoral forest to sea-level rise. Global Ecol. Biogeography 18, 98–110. doi:
10.1111/j.1466-8238.2008.00429.x
Virah-Sawmy, M., Gillson, L., Gardner, C. J., Anderson, A., Clark, G., and Haberle, S.
(2016). A landscape vulnerability framework for identifying integrated conservation
and adaptation pathways to climate change: the case of madagascar’s spiny forest.
Landscape Ecol. 31, 637–654.
Voarintsoa, N. R. G., Wang, L., Railsback, L. B., Brook, G. A., Liang, F., Cheng, H.,
et al. (2017). Multiple proxy analyses of a U/Th-dated stalagmite to reconstruct
paleoenvironmental changes in northwestern madagascar between 370 CE and 1300
CE. Palaeogeogr. Palaeoclimatology Palaeoecol. 469, 138–155.
Vorontsova, M. S., Besnard, G., Forest, F., Malakasi, P., Moat, J., Clayton, W. D., et al.
(2016). Madagascar's grasses and grasslands: anthropogenic or natural? Proc. R. Soc. B:
Biol. Sci. 283, 20152262. doi: 10.1098/rspb.2015.2262
Vorontsova, M. S., Lowry, P. P., Andriambololonera, S. R., Wilme, L., Rasolohery, A.,
Govaerts, R., et al. (2020). Inequality in plant diversity knowledge and unrecorded plant
extinctions: An example from the grasses of Madagascar. Plants People Planet. 3 (1),
45–60.
Waeber, P. O., Wilme, L., Mercier, J.-R., Camara, C., and Lowry, P. P. (2016). How
effective have thirty years of internationally driven conservation and development
efforts been in Madagascar? PLoS One 11, e0161115. doi: 10.1371/journal.pone.0161115
Watson, J. E. M., Whittaker, R. J., and Dawson, T. P. (2004). Avifaunal responses to
habitat fragmentation in the threatened littoral forests of south-eastern Madagascar. J.
Biogeography 31, 1791–1807. doi: 10.1111/j.1365-2699.2004.01142.x
Willis, K. J., Gillson, L., and Virah-Sawmy, M. (2008). Nature or nurture: the
ambiguity of C4 grasslands in Madagascar. J. Biogeography 35, 1741–1742.
doi: 10.1111/j.1365-2699.2008.01985.x
Wingard, G. L., Bernhardt, C. E., and Wachnicka, A. H. (2017). The role of
paleoecology in restoration and resource management—the past as a guide to future
decision-making: review and example from the Greater Everglades Ecosystem, USA.
Front. Ecol. Evol. 5, 11. doi: 10.3389/fevo.2017.00011
Wurz,A.,Tscharntke,T.,Martin,D.A.,Osen,K.,Rakotomalala,A.A.,
Raveloaritiana, E., et al. (2022). Win-win opportunities combining high yields with
high multi-taxa biodiversity in tropical agroforestry. Nat. Commun. 13, 4127.
doi: 10.1038/s41467-022-30866-8
Yang, Y., Tilman, D., Furey, G., and Lehman, C. (2019). Soil carbon sequestration
accelerated by restoration of grassland biodiversity. Nat. Commun. 10, 1–7.
doi: 10.1038/s41467-019-08636-w
Zaehringer, J. G., Eckert, S., and Messerli, P. (2015). Revealing regional deforestation
dynamics in North-Eastern Madagascar—Insights from multi-temporal land cover
change analysis. Land 4, 454–474. doi: 10.3390/land4020454
Gillson et al. 10.3389/fcosc.2023.1286459
Frontiers in Conservation Science frontiersin.org11
