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Estimating the Carbon Sequestration Ecosystem Services of the Savanna: the Case of the Bismarck Palm Bismarckia Nobilis Hildebr. &H.Wendl. in the Protected Area of Antrema, Madagascar

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Estimating the Carbon Sequestration Ecosystem

Services of the Savanna: the Case of the Bismarck Palm

Bismarckia Nobilis Hildebr. &H.Wendl. in the

Protected Area of Antrema, Madagascar

Mouna Chambon, Edmond Roger, Miadama Faramalala, Joro Rakotoarinivo,

Solofo Rakotondraompiana, Camélias Rasoamanantenaniaina, Tony

Ramihangihajason, Bernard Riéra

To cite this version:

Mouna Chambon, Edmond Roger, Miadama Faramalala, Joro Rakotoarinivo, Solofo Rakotondraom- piana, et al.. Estimating the Carbon Sequestration Ecosystem Services of the Savanna: the Case of the Bismarck Palm Bismarckia Nobilis Hildebr. &H.Wendl. in the Protected Area of Antrema, Madagascar. 2021. �hal-03101700�

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CHAMBON et al.

Estimating the Carbon Sequestration Ecosystem Services of the Savanna: the Case of the Bismarck Palm Bismarckia Nobilis Hildebr. &H.Wendl. in the Protected Area of Antrema, Madagascar

List of authors: Mouna CHAMBON, Edmond ROGER, Miadama FARAMALALA, Joro RAKOTOARINIVO, Solofo RAKOTONDRAOMPIANA, Camélias

RASOAMANANTENANIAINA, Tony ARISON, Bernard RIERA

Affiliations:

Mouna CHAMBON (corresponding author): UMR 7279 (Mecadev) CNRS et MNHN,

Département Ecologie, Gestion de la Biodiversité, Muséum national d’Histoire naturelle, 1 avenue du Petit Chateau, 91800 Brunoy, France.

✉mouna.chambon@gmail.com

Edmond ROGER: Département Biologie et Ecologie Végétales (DBEV),

Université d’Antananarivo, BP 566, Antananarivo 101, Madagascar

✉rogeredmond1@yahoo.fr

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Miadama FARAMALALA: Département Biologie et Ecologie Végétales (DBEV),

Université d’Antananarivo, BP 566, Antananarivo 101, Madagascar

✉ faramia21@gmail.com

Joro RAKOTOARINIVO: Institut et Observatoire de Géophysique d'Antananarivo (IOGA),

Université d’Antananarivo, BP 566, Antananarivo 101, Madagascar

✉ XXXXX

Solofo RAKOTONDRAOMPIANA: Institut et Observatoire de Géophysique d'Antananarivo

(IOGA), Université d’Antananarivo, BP 566, Antananarivo 101, Madagascar

✉srakotondraompiana@gmail.com

Camélias RASOAMANANTENANIAINA: Département Biologie et Ecologie Végétales

(DBEV), Université d’Antananarivo, BP 566, Antananarivo 101, Madagascar

✉ rasoamanantenaniaina@gmail.com

Tony ARISON: Institut et Observatoire de Géophysique d'Antananarivo (IOGA), Université d’Antananarivo, BP 566, Antananarivo 101, Madagascar

Bernard RIERA: UMR 7279 (Mecadev) CNRS et MNHN, Département Ecologie, Gestion de

la Biodiversité, Muséum national d’Histoire naturelle,

1 avenue du Petit Chateau, 91800 Brunoy, France.

✉bernard.riera@mnhn.fr

Received: _____________; Revised: ______________ (optional); Accepted _____________

ABSTRACTS

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Sustainable management of natural resources has become a critical issue in the context of climate change. Although savannas represent a significant carbon sink, they have long been neglected in carbon inventories. Here, we addressed the potential win-wins of biodiversity conservation and climate change mitigation in savanna through a particular focus on the endemic palm species Bismarckia nobilis Hildebr. &H.Wendl. in the New Protected Area (NPA) of Antrema, North West of Madagascar. The aim of the present study was to assess the sustainability of the current management of Bismarckia nobilis populations in the NPA by drawing up a prospective scenario and to determine the middle-term potential carbon sink strength of the savanna. We collected ecological data in six survey sites. Then, we quantified the carbon sequestration levels in

Bismarckia nobilis by using allometric relationships and spatial analysis. The data allowed us to build a model of change in Bismarckia nobilis population dynamics for the next 50 years. Our findings demonstrated the importance of carbon storage capacity of the species, with 90,000 tonnes of carbon currently stored in the total surface area of the savanna. Furthermore, we found that anthropogenic activities do not have a negative impact on the population size of the species in the NPA and rather, contribute to its expansion through regulated burnings. These results indicate the role of the savanna of the NPA as an important carbon sink in the middle term.

Keywords: Arecaceae, climate mitigation, conservation biology, nature-based solutions, participatory approach, sustainability, tropical climate

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La gestion durable des ressources naturelles est devenue un enjeu primordial dans le contexte du changement climatique. Bien que les savanes représentent un puit de carbone important, elles ont longtemps été négligées dans les inventaires de carbone. Ici, nous avons abordé la question des possibles co- bénéfices de la conservation de la biodiversité et de l’atténuation aux changements climatiques dans la savane au travers du cas particulier de l’espèce de palmier endémique, Bismarckia nobilis Hildebr. & H.Wendl., dans la Nouvelle Aire Protégée (NAP) d'Antrema au nord-ouest de Madagascar. L'objectif de la présente étude était d'évaluer la durabilité de la gestion actuelle des populations de Bismarckia nobilis dans la NAP et d’estimer le potentiel de la savane en tant que puit de carbone à moyen terme. Dans le cadre d’une mission de terrain, nous avons quantifié les niveaux de séquestration de carbone du palmier Bismarckia nobilis sur la base de relations allométriques et d’une analyse spatiale. Ces données nous ont permis de construire un modèle d’évolution de la dynamique des populations de Bismarckia nobilis pour les 50 prochaines années. Nos résultats montrent que l'espèce Bismarckia nobilis présente une capacité de stockage de carbone importante, estimée à 90 000 tonnes pour la surface totale de la savane à Antrema. En outre, notre modèle d’évolution prévoit une augmentation des populations de Bismarckia nobilis pour les 50 prochaines années. Les résultats de cette étude mettent en avant l’importance de la savane d’Antrema en matière de séquestration de carbone à moyen terme.

Mots clés: Approche participative, Arecaceae, atténuation du changement climatique, biologie de la conservation, climat tropical, durabilité, solutions fondées sur la nature

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1. INTRODUCTION

1.1. Literature review

1.1.1. Potential win-wins of biodiversity conservation and carbon storage incentives

In tropical areas, deforestation and forest degradation have reached alarming rates and cause substantial damages to biodiversity and its related resources (Food and Agriculture Organisation [FAO], 2016) (Table S1). Stopping deforestation is not only essential for biodiversity

conservation purpose, but also for climatic reasons given that the decrease in forest cover has a major impact on ecosystem services such as carbon storage (Olander et al., 2008). The Paris Agreement, which was adopted at the 21st Conference of the Parties (COP21) in 2015, reaffirmed the world's commitment to tackling climate change. The process of quantifying carbon

sequestration in terrestrial ecosystems has, therefore, emerged as a new tool to developing climate mitigation strategies (Vieilledent et al., 2012). Constituting two major crises of the contemporary epoch, also referred as the “anthropocene” (Crutzen, 2002), both climate change and biodiversity erosion are causing systematic changes all over the world. As a result, a synergic and integrated approach to tackling these two crises would achieve mutual co-benefits (Millenium Ecosystem Assessment [MEA], 2005).

1.1.2. Significance of savannas in the global carbon budget

Although savannas are acting as an important carbon sink, there is a gap in the literature on their role in climate change mitigation (Grace et al., 2006; Razanajatovo, 2013). Broadly speaking, savannas are defined as “any tropical ecosystem containing grasses, including woodland and grassland types” (Eiten, 1972), which encompasses a large range of savanna types. This biome covers one-sixth of the land surface, including 50% of the African continent (Campbell, 1996).

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Due to its large area of occupancy and combined with the frequency of burnings, savannas have a direct impact on the global carbon budget. It is estimated that up to 8 GtC of CO2 is emitted annually from savanna burning (Seiler & Crutzen, 1980). Furthermore, land-use change, through grazing and crop cultivation, contributes to increasing fire events and climate change is likely to increase the frequency of burning in the coming years (Grace et al., 2006). Therefore, managing savannas has emerged as a crucial concern for tackling climate change.

1.1.3 Participatory management of natural resources in Madagascar: case of Antrema NPA In Madagascar, savannas cover a substantial part of the total land surface, about 70%

(Razanajatovo, 2013). Madagascar has one of the greatest endemism rates in the world with notably 90% endemic vascular plants (Goodman & Benstead, 2003). However, this unique biodiversity is becoming more and more threatened (Blanc-Pamard & Ramiarantsoa, 2003). In 2003, at the Durban World Parks Congress in Durban, the Government of Madagascar made the commitment of tripling the surface area of its protected areas and launching a new Protected Area System (SAPM), which covers both the extension of the network of existing Protected Areas and the creation of New Protected Areas (NPAs) (Agence Française de Développement [AFD], 2015). Among these new protected areas, the NPA of Antrema is one of the first community protected area in Madagascar, which means that it is completely managed by a bottom-up logic (Pioch, 2015). In this NPA, woody savanna is dominated by the palm species Bismarckia nobilis Hildebr. & H.Wendl. (Dransfield & Beentje, 1995; Gauthier & Leclerc- Cassan, 2000). Palm species have been reported to be particularly relevant for assessing the resilience of ecosystems at a species level in tropical America (Montufar et al., 2011) (Table S2).

Here, we apply a similar approach by assessing the resilience of the savanna of the NPA and its climatic implications through the particular case of B. nobilis.

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1.2. Objectives

The present study aims to assess the sustainability of current management of B. nobilis

populations in the NPA of Antrema by drawing up a prospective scenario for the next 50 years and therefore conclude the middle-term potential carbon sink strength of the savanna ecosystem.

The specific objectives of this study were to: (a) quantify the carbon sequestration levels of B.

nobilis in the savanna of the NPA; (b) determine the key factors that affect the growth of the studied species and estimate their respective value.

1.3. Study site

The NPA of Antrema is located on the north-western coast of Madagascar (Figure S1). It covers 20, 620 ha of which 1, 000 ha is of the marine park. Throughout the year, temperatures remain high, with an average annual temperature of around 26°C. The annual average precipitation is quite high, with a value of 1, 498 mm per year. The station is characterised by two very distinct seasons: a warm and wet season (November to April) and a dry and fresh season (May to

October). The NPA of Antrema is part of the “Western Region” of Madagascar (Humbert, 1955).

It is included in the Western eco-floristic zone from 0 to 800 m altitude characterised by a climatic vegetation (Pioch, 2015).

2. METHODS

2.1. Study species

Madagascar is famous for its remarkably diverse palm flora with 171 species of palms

distributed in 16 genera and representative of 4 out of the 6 recognised subfamilies in the world (Dransfield et al., 2008). The majority of palm species in Madagascar occur in the rainforests of the Eastern and Northern part of the Island. However, there are four palms species, such as

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B. nobilis, that have adapted to secondary habitats like savannas (Dransfield & Rakotoarinivo, 2011). B. nobilis is a monotypic palm species, one of the three representatives of the subtribe Hyphaeninae of the Arecaceae family (Dransfield, 2006). It is a solitary, robust palm tree that is endemic to the Red Island. It is one of the most common palm trees in the north and west of Madagascar and occurs as the dominant tree species on savannas. Although it is widespread on the west coast of the country, little is known about its life-history (Dransfield & Rakotoarinivo, 2011). B. nobilis has the ability to survive fire, which explains its success in invading such anthropic habitats over time (Koechlin et al., 1974).

2.2. Fieldwork in the NPA of Antrema

During one month in the NPA of Antrema, we conducted an ecological survey in order to determine key environmental factors of the populations of B. nobilis in the area. The choice of the survey sites was defined based on field prospecting. A total of six survey sites were selected (Table 1). The Braun-Blanquet method was applied in each of the six survey sites (Braun-

Blanquet, 1964). In each site, two surveys were conducted, totalling 12 survey plots. Each survey plot respected three criteria: floristic homogeneity, structural and physiognomic homogeneity and uniformity of the ecological conditions. Each plot consisted of a 20 m × 50 m rectangle set perpendicular to sea level. They were then subdivided by fluorescent flags into ten parcels of 10 m × 10 m. The delineation of the plots and their subdivisions was done with string. Within each plot, we recorded the following environmental factors for all B. nobilis individuals of a height greater than 10 cm: Diameter at Breast Height (DBH) with Breast Height set at 130 cm, maximum height, phenological state and number of leaves. The DBH was measured along the stipe using tape measure. We estimated the height based on trigonometric methods using an Abney’s level (Philip, 1994). Finally, we classified each recorded individual according to its

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development state (Table S3). The geographical location of each plot was obtained by using a GPS and the topography with a clinometer

2.3 Analysis of data 2.3.1 Spatial analysis

In this study, we used a moderate spatial resolution image (15 m) from the Landsat 8 satellite (launched February 11, 2013) from the U.S Geological Survey. This image was acquired over the NPA of Antrema in March 2017 and comprised 11 spectral bands in total. We pre-processed the satellite image (virtual raster building; colour calibration; polygon clipping; area

delimitation) and then applied a supervised classification involving seven Region of Interests (ROIs). All analysis was performed on QGIS software (version 2.14.3-Essen). The classification output enabled us to conduct further analysis based on the estimation of the surface area of the class of savanna with B. nobilis.

2.3.2 Quantifying carbon sequestration in Bismarckia nobilis

For each survey site, we quantified the above-ground carbon of B. nobilis using allometric relationships. Two equations were explored in order to first determine the above-ground biomass (AGB) of B. nobilis (Table S4). Equation 1 is an equation proper to the study species

(Razanajatovo, 2013), whereas equation 2 is employed as a general equation for the Sabal genus (Delaney et al., 1999; Brown et al., 2001). These two equations express the AGB as a function of the maximal height of individuals. Comparison of results based on these two equations is

expected to confirm the relevance of using specific allometric equations. In this study, we did not take account the herbaceous biomass because we assumed that changes in carbon pools are greater in palms than herbaceous species (Pellegrini et al., 2016). Finally, above-ground carbon per site was deduced from the AGB values, assuming that biomass is 50% carbon (Chave et al.,

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2005). Total carbon stocks per site were obtained by summing up the values of carbon stocks of the three following compartments: aboveground biomass, roots, and litter. Owing to time and logistic constraints, we could solely measure above-ground biomass, so we used estimations from previous studies for roots (1.9-6

t/ha) and litter (7.13-4

t/ha) (Razanajatovo, 2013). Finally, the overall potential for carbon sequestration of B. nobilis in the NPA of Antrema was calculated on the basis of the previous spatial analysis by multiplying the total surface area of the savanna with the average carbon stock of the species per hectare.

2.3.3 Prospective model of Bismarckia nobilis population dynamics in the NPA

We created a prospective model of the population dynamics of B. nobilis in the NPA using a linear regression. First, we compiled data on the excel software (version 15.2) and subsequently applied the equation below (Equation 3). We considered the six following factors: the current population size of B. nobilis in the NPA (Pn); the annual mortality rates of the species related to roofing (m1), handicraft activities (m2), and fire (m3); the natural mortality (m4) and

regeneration (r) rate of the species. The current population size of the species was estimated by multiplying the average palm density per hectare (564 indiv /ha) with the total surface area of the savanna. In addition, we used results from a complementary study on the local uses of B.nobilis in the NPA, which estimates that 87, 333 palm individuals are harvested per year for roofing and 700,000 palms for handicrafts. The reiteration period covers 50 years. The final model comprises three prospective scenarios that differ in the estimation of the mortality rates as given in Table 2.

Equation 3: Y = Pn - (87,333×m1 + 700,000×m2 + Pn×m3 + Pn×m4) + Pn× r

Where: Pn is the population size of B. nobilis at t 0 Y is the population size of B. nobilis at t+1 (Pn+1)

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3. RESULTS

3.1 Carbon stocks by site

The average carbon stocks by survey site according to the two allometric equation types are summarised in Table 3. These results show that the average values are similar in the two cases.

Results based on the general equation for the Sabal genus show slightly higher values (X= 13.6 +/- 7 SD t/ha) than those based on the equation proper to the species B. nobilis (X=12.215 +/- 6 SD t/ha). But both cases indicate the same general distribution of carbon stocks between sites.

The highest average value is observed in North-West of Sahariaka Lake with 21.3 t/ha (Equation 1) and 24.4 t/ha (Equation 2) (Figure 1). By contrast, the sites of South-east and South –west Antrema are characterised by the lowest average values: respectively ranging from 4.5 t/ha to 4.71 t/ha and from 6.39 t/ha to 6.89 t/ha.

3.2 Carbon storage in the palm savanna

Results from the spatial analysis are displayed on the map of the NPA below (Figure 2), which has been classified in seven categories. For the purpose of this study, we focused on the class

“Savanna and palm” only, without considerations to the other classes which might be a matter of further discussions. The total surface area of the palm savanna is valued at 7,417.4 ha out of the 20, 620 ha of the NPA, which represents 35.9 % of the total area. Based on this calculation and taking account the species density per survey site (Table S5), we estimated the total population size of B. nobilis in the palm savanna of the NPA 4 million individuals, as showed in Table 4.

Finally, the total value of carbon stored in B. nobilis in the savanna of the NPA is higher according to the equation for the Sabal genus than if we use the equation proper to the target species: respectively 100, 877 and 90, 604 t of carbon.

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3.3. A prospective study on the sustainable dimension of the exploitation of Bismarckia nobilis.

The comparison of the three scenarios are illustrated on Figure 3. None of these three scenarios indicate a total depletion of populations over the next 50 years. However, scenarios n°1 and n°2 show a significant degradation of the resource, whereas scenario n°3 rather suggests an increase of populations. Scenario n°1 points to a rapid and substantial decrease in the population size of B. nobilis. It indicates a drop of 60% of populations by 2067: from 4 million to 1.6 million of individuals. Although less critical, scenario n°2 also indicates a drop of the population size of 22% by 2067. By contrast, scenario n°3 shows an extensive increase in the population size. From 4 million, we reach almost 6 millions of individuals in 2067.

4. DISCUSSION

4.1 Potential carbon sink strength of the savanna with B. nobilis in the NPA

In this study, we applied two allometric equations for estimating the carbon sequestration potential of the savanna in the NPA. In the two cases, our results show similar values. We preferentially chose to base our interpretations on the first equation, since it is for B. nobilis specifically and reflects a higher accuracy. In this study, we estimated the value of total carbon stocks of B. nobilis in the savanna at 12.215 t/ha and demonstrated the variability in terms of carbon stocks among the six survey sites. Carbon stocks in North-West Sahariaka Lake are the highest, whereas the lowest carbon stocks are found in South-east and South-west Antrema.

These differences may be related to palm density in each site since there is a direct link between biomass accumulation and carbon stocks (Nizami et al., 2017). Regarding the outcome of our spatial analysis, we estimated the total surface area of the savanna to be 7,417.4 ha, which

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represents 35.9 % of the NPA. These findings are congruent with a previous research, which estimated a value of 35. 78 % of the NPA (Razanajatovo, 2013). In complementing field measurements, spatial analysis turned out to be of a great utility for estimating carbon

sequestration levels in B. nobilis within the NPA. A better outcome could have been produced with higher resolution satellite imagery, but this methodological choice would have exceeded our budget. As reported in several academic studies, satellite data now has a great potential for identifying co-benefits between biodiversity conservation and carbon storage (Olander et al., 2008; Hansen et al., 2013; Pfeifer et al., 2016). According to similar studies that were carried out on other vegetal formation, B. nobilis savanna shows an intermediate carbon storage capacity, lower than pristine forest, which is of utmost importance in terms of carbon sequestration, but much higher than herbaceous savanna (Table S6). Although falling in the same order of

magnitude, our findings are slightly lower than that of a previous research that was conducted in the savanna of the NPA as well (Razanajatovo, 2013). The fact that we did not include the carbon stocks of herbaceous species in the savanna, but focused on the B. nobilis species only, might explain this contrast. Moreover, our calculations do not include the carbon amount that is stored in palm leaves and the leaves contain about 31.7% of the total biomass in the case of the palm species Attalea speciosa Mart. (Gehring et al., 2011). By taking account the leaf biomass of B. nobilis, we could therefore have increased its carbon storage capacity.

Overall, the species B. nobilis shows a great carbon storage capacity since its total carbon stock in the savanna is about 90,604 t without considering the leaf biomass. Comparing to previous studies that were conducted in the NPA, this storage capacity is lower than that of the dry dense forests but represents double of that of the mangrove formation (Table 5). Therefore, the

abundance of B. nobilis in the savanna of the NPA makes it a critical carbon sink in the long

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term. Although savannas are not a priority ecosystem in REDD+ mechanisms, this study falls within a substantial academic corpus that shows their potential in terms of carbon storage (Grace et al., 2006; Pelligrini et al., 2016). According to Scurlock & Hall (1998), the global carbon sink strength of savannas might even be valued up to 0.5Gt. Besides biomass accumulation, the carbon storage capacity of savannas is influenced by the frequency of fires and grazing worldwide. Therefore, protecting savannas from anthropogenic pressures would considerably increase their carbon sink potential (Grace et al., 2006).

4.2 Middle-term availability of the studied natural resource

In this study, we tested different estimations of mortality rates of B. nobilis related to fire, roofing, and handicrafts activities as well as its natural mortality and regeneration rate.

According to these estimations, we came up with three prospective scenarios on the availability of the resource over the next 50 years. Scenario n°1 points to a rapid and substantial decrease in the population size of B. nobilis. This result reflects that the natural regeneration rate is too low to catch up with mortality rate related to anthropogenic activities. By contrast, scenario n°3 shows an extensive increase in the population size. In this situation, the related natural regeneration rate is two times higher than in scenario n°1 and exceeds the losses due to anthropogenic activities. Finally, scenario n°2 is an intermediate scenario that excludes fire events but considers a mortality rate from roofing activities higher than the two other scenarios, whereas the natural regeneration rate is lowered to 1% only. As a result of this disequilibrium, the scenario shows a significant decrease of the population size. Natural regeneration and anthropogenic activities, which include fire and local uses, seem to act as two driving forces on the population dynamics of B. nobilis species. Based on evidence from the field assessment and local perceptions, we conclude that scenario n°3 is the most appropriate scenario to describe the

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current situation within the NPA. On the middle-term, the overall population of Antrema is not under a risk of depletion but will rather experience an extension phase. Anthropogenic activities do not have a negative impact on the population size and rather contribute to its expansion through fire that stimulates seed germination. Given the substantial carbon storage capacity of B.

nobilis, as stated previously, we can cautiously predict an increase in the carbon sink potential of the savanna of the NPA over the next 50 years. These findings contrast with two previous studies that were carried out in other parts of Madagascar (Table S7). Their conclusions indicate a severe depletion of B. nobilis populations, either on the short–term (Ratoavimbahoaka, 2006) or the middle-term (Rabefarihy, 2007). These assessments differ from that of our study because the two studied areas are not part of any protection scheme like in the case of Antrema, which is a

category VI Protected Area according to the IUCN classification (Pioch, 2015). Moreover, the cultural usages that characterise these regions are associated with a more intensive exploitation of the species (Table S7). These two alarming scenarios do stress the necessity to continue monitoring the population dynamics of B. nobilis because some threats are common to all regions, and therefore likely to challenge populations in Antrema in the long-term.

5. CONCLUSION

This research contributed to highlight the significance of savannas in carbon sequestration. In this case study, we found that the average value of carbon that is stored in B. nobilis in the savanna of Antrema is 12.215 t/ha, based on an allometric equation proper to the species. Given that the total surface area of the savanna was estimated at 7, 417 ha, which represents 35.9% of the NPA, we estimated an amount of 90, 604 t of carbon stored in B. nobilis in the study area, without considering its leaf biomass. Although lower than the carbon sequestration levels in the

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dry dense forests, this value represents double of that of the mangrove formation in the NPA (Rakotovao, 2013). Furthermore, B. nobilis turned out to be a good indicator of the resilience of the savanna ecosystem at a species level. Scenario n°3 was chosen as the most appropriate scenario for describing changes in the palm population over the next 50 years. Natural

regeneration and anthropogenic activities, which include fire and local uses, seem to act as two driving forces on population dynamics of the B. nobilis species. Anthropogenic activities do not have a negative impact on the overall population but, rather, contribute to its expansion through fire that stimulates seed germination. In the middle term the resource is not under risk of

depletion but will rather experience an extension phase. Therefore, our findings highlight the mutual co-benefits of sustainable use of natural resources and climate change mitigation.

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TABLES

Table 1. Location of the six survey sites in Antrema NPA with their GPS coordinates Location of the site GPS coordinates

North-west of Sahariaka Lake Plot 1: 15° 44,658' S, 046° 07,492' E Plot 2: 15° 44,698' S, 046° 07,312' E West Kapahazo Plot 3: 15° 45,675' S, 046° 06,634' E Plot 4: 15° 45,696' S, 046° 06,666' E

Ankokoala Plot 5: 15° 45, 974' S, 046° 09, 504' E

Plot 6: 15° 45, 917' S, 046° 09, 538' E Maskoamena Plot 7: 15° 45, 623' S, 046° 10, 621' E Plot 8 :15° 45, 606' S, 046° 10, 656' E South-east Antrema Plot 9: 15° 43, 400' S, 046° 10, 350' E Plot 10:15° 43, 442' S, 046° 10, 341' E South-west Antrema Plot 11:15° 43, 126' S, 046° 09, 738' E Plot 12:15° 43, 099' S, 046° 09, 813' E

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Table 2. Mortality and regeneration rates (%) applied to the species Bismarckia nobilis according to the three scenarios of this prospective study

Rates (%) Scenario n°1 Scenario n°2 Scenario n°3

Mortality related to roofing (m1) 2 5 2

Mortality related to harvest (m2) 1 2 1

Mortality related to fire (m3) 2 0 2

Natural mortality (m4) 1 1 1

Regeneration (r) 2 1 4

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Table 3. Biomass and carbon stocks (t/ha) by survey site according to the two allometric equations. Equation 1 (AGB = 12.72H + 12.90) is proper to Bismarckia nobilis (Razanajatovo, 2013) and Equation 2 (AGB = 24.553 + 4.921H + 1.017H2) is generally used for the Sabal genus (Delaney et al., 1999; Brown et al., 2001)

Survey site Biomass Equation 1 (t/ha)

Carbon stocks Equation 1 (t/ha)

Biomass Equation 2 (t/ha)

Carbon stocks Equation 2 (t/ha) North-west of

Sahariaka Lake

42.6 21.3 48.8 24.4

West Kapahazo 26.2 13.1 30.8 15.4

Ankokoala 29.8 14.9 33.2 16.6

Maskoamena 26.2 13.1 27.4 13.7

South-east Antrema

9 4.5 9.42 4.71

South-west Antrema

12.78 6.39 13.78 6.89

Mean 24.430 (+/- 6 SD) 12.215 (+/- 6 SD) 27.2(+/- 7SD) 13.6 (+/- 7SD)

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Table 4. Population size of Bismarckia. nobilis in the NPA of Antrema and related carbon stocks for the total surface area of the savanna, calculated based on the classification output resulting from the spatial analysis

Average density in

B.nobilis

(average number of individuals/ha)

Surface area of the palm savanna (ha)

Total population size in the NPA (number of individuals)

Total carbon stocks of B.nobilis in the palm

savanna (t)

564 7,417.4 4,183,413 ~ 4

million

90,604 (Eq.1)

100,877 (Eq.2)

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Table 5. Comparison of carbon stock estimates in different vegetal formations of the NPA Reference Rasoamanantenaniaina (2016) Chambon (2017) Rakotovao (2013) Vegetal

formation

Dry dense forest Savanna with

B.nobilis

Mangroves

Surface area (ha) 4,379 7,417.4 1,764.46

Carbon stock (t/ha)

54.6 12.215 5.471

Total carbon stock (t)

115,182 90,604 9,653

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FIGURE LEGENDS

Figure 1. Carbon stocks by survey site according to the two allometric equations.

Figure 2. Classified map of the New Protected Area of Antrema. We used a satellite image from Landsat 8 (2017) with moderate spectral resolution (15m). The image was processed on QGIS (Version 2.14.3-Essen).

Figure 3. Changes in the population size of Bismarckia nobilis in the NPA over the next 50 years according to the three scenarios, as defined in Table 2.

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FIGURES Figure 1

0 5 10 15 20 25 30

North-West of Sahariaka

lake

West Kapahazo

Ankokoala Maskoamena South-East Antrema

South-West Antrema

Carbon stocks (t/ha)

Villages

Equation 1 Equation 2

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Figure 2

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Figure 3

0 1000000 2000000 3000000 4000000 5000000 6000000 7000000

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30 32 34 36 38 40 42 44 46 48 50

Population size (number of individuals)

Years

scenario 1 scenario2 scenario3

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ACKNOWLEDGEMENTS

I would first like to thank my thesis supervisor Dr. Bernard Riéra of the Adaptive Mechanisms and Evolution department (UMR 7179) at the National Museum of Natural History of Paris. His help was precious throughout this research project. I would also like to thank my thesis co- supervisor Dr. Edmond Roger and Mrs. Amelie Faramalala of the Department of Ecology and Vegetal Biology at the University of Antananarivo. They were deeply involved during my time in Madagascar and my fieldwork could not have been successfully conducted without their inputs. I would also like to acknowledge Dr. Farid Dahdouh-Guebas of the Systems Ecology and Resource Management Department at ULB and Dr. Claude Marcel Hladik of the Man and Environment department (UMR 7206) at the National Museum of Natural History of Paris, as my two examiners. I am also gratefully indebted to the experts who have guided my work analysis with patience and strong support: Dr. Flora Pennec of the UMR 7206 at the National Museum of Natural History of Paris and Anoumou Kemavo, GIS and Forest Expert Project Officer at the National Agency of Forests (ONF). In addition, I would never have been able to conduct my fieldwork and collect my data without the support and guidance of my assistant Camelias Rasoamanantenaniaina. Many thanks to all people I met in Madagascar and who participated to my overall understanding of the region and research context.

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