# Coexist or disappear: An interdisciplinary and longitudinal study of the relationship between humans and lemurs on the island of Mayotte

Claire Harpet, Laurent Tarnaud, Bernard Riéra, Bruno Simmen

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Claire Harpet, Laurent Tarnaud, Bernard Riéra, Bruno Simmen. Coexist or disappear: An inter-disciplinary and longitudinal study of the relationship between humans and lemurs on the island of Mayotte. Revue de Primatologie, 2025, 15, pp.24117. hal-05377197

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# Coexist or disappear: An interdisciplinary and longitudinal study of the relationship between humans and lemurs on the island of Mayotte

*Coexister ou disparaître : Étude interdisciplinaire et longitudinale des relations entre humains et lémuriens sur l'île de Mayotte*

Claire Harpet, Laurent Tarnaud, Bernard Riéra and Bruno Simmen

<figure><img src="image_2.png" /></figure>

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URL: https://journals.openedition.org/primatologie/31766

ISSN: 2077-3757

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Société francophone de primatologie
# Coexist or disappear: An interdisciplinary and longitudinal study of the relationship between humans and lemurs on the island of Mayotte

*Coexister ou disparaître : Étude interdisciplinaire et longitudinale des relations entre humains et lémuriens sur l'île de Mayotte*

Claire Harpet, Laurent Tarnaud, Bernard Riéra and Bruno Simmen

## EDITOR'S NOTE

Received 13/05/2024, accepted after revisions 12/12/2024, published online 01/04/2025 in the context of the special issue: 'Conservation' and 'Coexistence' in primatology: antagonisms and overlaps (2024).

This version is the English translation of the article recorded under the following reference:

Harpet C, Tarnaud L, Riéra B, Simmen B. 2024. Coexister ou disparaître : Étude interdisciplinaire et longitudinale des relations entre humains et lémuriens sur l'île de Mayotte. Revue de primatologie 15.

This article was written before Cyclone Chido struck Mayotte on December 14th 2024.

## 1 Introduction

"It's like we'd levelled a promontory, knocked down a mountain - the world suddenly seemed to have shrunk, and now we're going to have to scrap with each
other to even exist in this corridor. It's like a whole swathe of reality that's collapsing, a complete and articulated conception of phenomena that our philosophy will lack from now on" (Eric Chevillard, 2005).

1. On the island of Mayotte, one of the four islands that make up the Comoros Archipelago that became a French department in 2011, the existence of the species *Eulemur fulvus* and humans is very closely intertwined. The first human populations to arrive on the island from Madagascar in the 8th-9th century introduced the *ankomba* or *komba* - from its vernacular name in Shibushi and Shimaore, respectively the Malagasy and Comorian languages of Mayotte (Allibert *et al.*, 1989). The lemur is more commonly known as 'the maki' and is a valued protected species in local culture because of the mythology of its origins and its more recent popularity with tourists. However, makis have moved closer to areas previously occupied by humans due to the transformation of Mayotte's economy from subsistence to cash, coupled with the rapid growth of the human population and the resulting increased pressure on forest environments. This has changed the relationship between humans and makis. Today, Mayotte's forest cover is mainly made up of converted forests and a few remnants of primary forests on the ridges. The latter have recently been classified as protected areas and cover around 8% of the territory (https://reserves-naturelles.org/reserves/forets-de-mayotte/). In the second half of the twentieth century, the expansion of agricultural activity and the development of infrastructure led to the island losing 20-30% of its forest areas (Lainé *et al.*, 2010). The gradual disappearance of the makis' habitat means their fate is even more closely linked to that of humans, in a local context of worrying socio-political, cultural and environmental crisis. Molecular and anatomical studies have found that the population of brown makis in Mayotte is close enough to the populations of *Eulemur fulvus* in north-west Madagascar to justify the use of the *E. fulvus* nomenclature (Mittermeier *et al.*, 1994; Pastorini *et al.*, 2000; Groves, 2001). However, Mittermeier *et al.* (1994) also found that hybrids involving another species of *Eulemur* (possibly *E. rufus*) cannot be ruled out.

2. It is important to take into account the ecology and demography of the Mayotte brown maki to understand current tensions. It is a gregarious and primarily frugivorous species which also includes substantial proportions of leaves in its diet, mainly during the dry season. Over 90 plant species that belong to 43 botanical families are eaten by *Eulemur fulvus* in Mayotte. This list is based on direct observations and includes all of Mayotte's woody fruit species that are exploited by humans (Tarnaud, unpublished). This species is found in all the forest types on Mayotte (mesophilous and hyrophilous evergreen forests at altitude and dry to semi-dry deciduous forests) along with small forest areas of less than one hectare that have been heavily modified by human intervention (Figure 1). An interdisciplinary team of primatologists, ecologists, botanists and anthropologists has been observing and analysing changes in ecosystems and human/lemur interactions on Mayotte's territory (insert no. 1) for the last twenty-five years (1999-2024). The latest work on demographic monitoring of the brown makis, coupled with observations and semi-directed interviews with farmers, was carried out in 2021-2022. This confirmed the changes that had already been observed in 2000 and was reinforced by similar monitoring in 2010 and 2015: (i) the maki population is continuing to decline (Tarnaud, 2024); (ii) local representations and beliefs about the maki are gradually losing their protective influence because of the nuisance this fruit-eating animal can cause in agroforests (Harpet, 2011); (iii) changes in representations of
the maki go hand in hand with changes in Mayotte's agricultural model and the status
of farmers there.

Insert 1

- 1999-2002: **ECOFOR project (Forest ecosystems): Inter-species interactions in Mayotte, variations in biodiversity and perceived heritage values.**

- 2010: *An eco-anthropological study of farmer/lemur relations in Mayotte - diagnosis, issues, recommendations*, DREAL Mayotte.

- 2010-2012: **Demographic trends in the Mayotte brown lemur population from 2010 to 2012, DEAL Mayotte** (Convention N°2010/DAF/SEF et N°2011/N°2012/DEAL/SEPR).

- 2015: *The Mahoran landscapes – an anthropological view*, DEAL Mayotte.

- 2016: *A census of brown lemurs on the island of Mayotte*, DEAL Mayotte (convention N°2016 /DEAL/SEPR).

- 2016-2019: **Census in agricultural areas of lemurs on the island of Mayotte**, Conseil Général (Convention N°DEDDE/DGA PATDD/CD/2016).

- 2018: **Study on the harvesting of wildlife - brown lemurs and fruit bats - on fruit production in the agrosystems of Mayotte**, CAPAM-ODEADOM (convention N°2015-23/avenant N°3).

- 2021-2023: 'The brown lemur in Mayotte, an emblematic protected primate that has become a pest? Eco-biology, perception/ representation, and food security in Mayotte'. Project funded by the François Sommer Foundation.

The authors' research missions in Mayotte.

<footnote n="3">3 In this article, we propose to retrace the main details of the evolution of the place of lemurs on the island of Mayotte in proximity to and with humans. This description begins with the first eco-anthropological studies from the end of the last century and continues up to the present day. The aim is to gain a better understanding of the challenges of human/lemur coexistence in a context of strong anthropic pressure and climate crisis.</footnote>
<figure><img src="image_3.png" /><figcaption>Figure 1</figcaption></figure>

Brown makis eating a mango (Photo credit: Laurent Tarnaud)

## 2 Methodology for a longitudinal interdisciplinary research study

The specific feature of our research is the monitoring (through occasional but regular field missions) of the relationship between the brown maki and human populations on the island of Mayotte. This article reviews an interdisciplinary approach applied to the study of cultural and biological dynamics and changes on the island and, more specifically, of how the coexistence of two species, humans and maki, has evolved. At the end of the twentieth century, the realisation that little was actually known about Mayotte's flora and fauna led the scientific community from France's National Museum of Natural History (MNHN) to organise targeted census programmes on the island. The first field mission combined taxonomic, ecological and anthropological approaches, the aim being for interdisciplinary researchers to work together to compile inventories of fauna and flora, observe the stability of biological populations and gather information on the viewpoints and practices of local populations. To successfully carry out the research programme and attempt to shift from a multi-disciplinary approach (each discipline carries out its research individually with the results subsequently aggregated) to an inter-disciplinary approach, the usual practice for the project was for fieldwork to be carried out jointly and to work on participatory observations (a common practice in the humanities and social sciences) involving team members. For this, all the researchers take an interest in work from other disciplines, pool skills and discuss their fieldwork together in situ. This is a demanding method that has been facilitated by the long-term involvement over 25 years of the same team of researchers. The results are based on joint contributions from disciplines in the life sciences (primatology, ecology, botany) and the humanities and social sciences (history and anthropology) as a result of a collective process.

### 2.1 The demography of the brown maki and eco-ethology

To document the demographic evolution of brown makis on the main island of Grande Terre since 1999, the line transect method was chosen for its robustness. This method is
classically used by ecology researchers to survey animal populations, particularly primates (Marshall *et al.*, 2008). Selective censuses were carried out across all of Grande Terre's forested and cultivated areas (Table 1). The density estimates were carried out on an ad hoc basis (Table 1) by measuring the perpendicular distance separating the centre of the group of brown makis from the counting line (Whitesides *et al.*, 1988). These measurements were then related to the surface areas occupied by the island's different natural or transformed ecological zones on the basis of the available IGN maps (Figure 2). Distance software (v6.0, release 2) was used to carry out the density calculations. We considered independently the specific case of makis captured by private individuals or from wildlife trafficking before being transferred to the nearby islet of Mbouziy.

<table><caption>Table 1</caption><thead><tr><td>Period</td><td>Number of lines of counting</td><td>Number of repetitions of counting</td><td>Length of counting lines (kms)</td><td>Season</td><td>Number of observers</td><td>Environment</td></tr></thead><tbody><tr><td>1999</td><td>17</td><td>1</td><td>[2.75-9]</td><td>Dry</td><td>1</td><td>Forest reserves and cultivated areas</td></tr><tr><td>2008</td><td>14</td><td>3</td><td>[0.5-1]</td><td>Dry</td><td>1</td><td>Forest reserves and cultivated areas</td></tr><tr><td>2010 à 2012</td><td>11</td><td>4</td><td>[0.66-1]</td><td>Dry</td><td>1</td><td>Forest reserves</td></tr><tr><td>2016</td><td>9</td><td>4</td><td>[0.66-1]</td><td>Dry</td><td>1</td><td>Forest reserves</td></tr><tr><td>2016 à 2019</td><td>4</td><td>4</td><td>[1-1.7]</td><td>Dry and wet</td><td>3 to 4</td><td>Agroforestry areas</td></tr></tbody></table>

Census of brown makis on Grande Terre. The census periods are indicated for each environment sampled along with the number of counting lines and repetitions for each of these lines. The minimum and maximum lengths of the counting lines are also given.

6

In 2021-2022, the centre of the island was the favoured geographical study area for an in-depth mirror study of ecological and anthropological data. The operational mode for our new fieldwork differed between the 2002 and 2010 studies, particularly with regard to the primatological aspect, because it focused specifically on the activity of makis in anthropised areas. The eco-ethological analysis was thus carried out on two areas of cultivated agroforest in the centre of the island (Figure 2). The first plot at Tsararano is used to cultivate cash crops while the second at Kahani is part of a food production system (see Figure 2). The Tsararano plot is a polyculture essentially composed of zones of cultivated and indigenous trees (mango, guava, cytherea plum, carambola, avocado, yoursop, *Albizia spp.*, breadfruit, jackfruit, etc.) and small banana plantations interspersed with more open areas where passion fruit, pineapple and papaya are grown. The same species, together with citrus fruits, are generally found in the second plot, which is much less structured in terms of identifiable areas because of the main crops grown therein. The 'instantaneous and scan sampling' method (with 5-minute intervals between scans) was used to determine the brown makis' activity budget and diet with distinctions made between the different parts of plants they eat and the maturity of the plant parts (Altmann, 1974). Between March and early July 2022, four and two focal groups respectively were tracked on each of the plots and GPS readings were taken during their movements to delimit the areas they used. The animals became
used to the observers straight away or after a few days of contact, and the groups,
which were cohesive wholes, were then continuously monitored from dawn to dusk or
by half-day, not including interruptions due to the weather (total: around 300 hours
spread over 28 days between the two sites, N = 3410 scans). In 2016, an estimate was
also made of fruit harvesting on the Tsararano plot by brown makis and in a banana
plantation to the north-west of the village of Tsingoni. The same protocol was used to
observe the animals' activities.

## 2.2 Ethnographic research and methods

7 In parallel to the census of brown makis, ethnographic surveys were also carried out more specifically among the farming community (farmers, government structures, institutions, associations) and also younger children and teenagers from Mayotte. Ethnographic research is based on a corpus of data gradually constructed as the missions progressed using qualitative methods (observation, in-depth semi-structured interviews, life stories, artistic productions). As is traditional in the field of ethnography, analysis is enriched by each field survey, based on an inductive approach focusing on observed situations and the lived realities of the survey respondents. The semi-structured interview was the preferred method for the project. This differs from a closed questionnaire and is of course carried out in situ. This method of collecting data is not restricted to pre-established questions and instead is gradually constructed as the discussions progress. It is based on a thematic grid that acts as a guiding framework without being reductive (see below). The interview consisted of spontaneous exchanges with any forms of digression accepted. The starting point was an informal conversation to avoid the discussion being directed according to the researcher's expectations. There was no time limit on the survey as the respondent had the choice of when to end a conversation. The data was recorded on a Dictaphone with the respondents' permission and then recorded in a field notebook. The interviews took place in the homes of local residents or in the agroforestry fields (the study site) particularly during participatory observations such as visits, festivities, taking part in harvesting, surveillance and all other daily activities in the fields.

Insert 2

Standard thematic grid for an ethnographic survey of the relations between
humans and brown maki:

- Knowledge of makis (ecology, behaviour, diet);

- The frequency of makis on agroforestry plots;

- Stories from daily life, knowledge, representations, anecdotes about makis and their relationship with humans;

- Changes observed (from makis to the socio-ecosystem).

8 The ethnographic study covered the whole island and took its specific linguistic and cultural characteristics into account.
9 French is the institutional and official language since its imposition at the time of colonisation. It has also developed through the schooling process, but two native languages are still spoken in Mayotte :

- Shimaore is a language of Bantu origin that is fairly close to the languages of the other three Comoros islands and is spoken in the centre and north of the island,

- Shibushi is spoken in the south of the island in around ten villages and is a language of Madagascan origin (Gueunier, 2016). Along with archaeological evidence this confirms the importance of the presence of Madagascan populations which were established very early on Mayotte's coasts (Allibert et al., 1989; Jaquin, 2009). Shibushi - also written as Kibushi - is the only Malagasy language spoken outside of Madagascar itself. This linguistic distribution has barely changed since the French settlers arrived in the 19th century.

10 In addition to this information about the two local languages, it should be noted that the Muslim religion taught in Koranic schools and Mosques strongly influences daily life on the island. The process of Islamisation of Mayotte started as early as the 12th century but only reached the entire population of the island around the end of the nineteenth century (Blanchy, 1990) when it was grafted onto the practice of Madagascan-influenced ancestor worship in the south of the island. These two belief systems are, and remain, decisive factors if one wishes to understand the representations and relationships developed by human societies with local biodiversity in Mayotte, and more specifically with the maki.

11 In the 25 years since then, whilst the selected respondents may have changed over the course of the research missions, the main composition of the sample groups (30 or more people per sample) stayed the same - men and women of all generations encountered in the fields and/or during social contacts during the researchers' immersion in the villages. It was never difficult to find a sample of farmers because this is of course a common or even ordinary activity in Mayotte. To work with children and teenagers (5-19 years) we relied on the educational structures that agreed to integrate our project into their school curriculum. These were primary schools in Mronabeja, a general secondary school in Sada, an agricultural secondary school in Coconi and the University of Mayotte which opened in the commune of Dembeni in 2011.

12 The first interviews were carried out between 2000 and 2002 following on from the transects the primatologists created to count and monitor makis crossing the agroforestry plots concerned (Tarnaud & Simmen, 2002).

13 In parallel with the surveys carried out along the transects, the researchers benefitted from immersion visits involving actually living with local people in villages covering three geographical areas:

- in the south of the island: the villages of Mronabeja and Moutsamoudou;

- in the north of the island: M'tsapéré;

- in the centre of the island: Coconi which the successive interdisciplinary research teams between 2000 and 2010 also used as their base camps.

14 In addition to these interviews, in 2002 a pictorial study was carried out with children from local nursery and primary schools to assess their knowledge of and sensitivity to makis in a context of growing land ownership of the forest (Harpet & Pagezy, 2010)

15 In 2010, the interviews were carried out during field visits to farmers organised by the Brigade Nature de Mayotte and based on information gathered ten years earlier, the aim
being to identify changes and, conversely, constants in Mayotte farmers'
representations and practices as regards makis. The farmers interviewed in 2002 were
either contacted again or found directly in their fields while new respondents were
identified, particularly young farmers. The interview locations were selected according
to the same criteria as in 2002, namely: (i) by following the transects carried out on the
periphery of agricultural plots for the 'census of Mayotte brown lemurs' project (Figure
2. This approach meant we could directly target farmers whose crops were on the edges
of forested areas); (ii) in villages that are representative of the selected communes - a
first sector in the north in the communes of Acoua, M'tsamboro and M'tsamgamouji,
and more specifically in the villages of M'tsamboro, M'tsamgamoudji and
Chembényumba; a second sector in the centre in the communes of Tsigoni and
Ouangani, and more specifically in the villages of Sohoa, Coconi, Ongojou, Kwalé and
Hajangoua; a third sector in the south in the commune of Kani-Keli, and more
specifically in the villages of Mronabeja, Passi Keli and Moutsamoudou.

16

A research mission in 2015 led to additional observations being made on the relations
between humans and makis in agroforestry environments, in the framework of more
specific work on landscapes (Harpet, 2016).

17

Finally, in 2021-22, collaborative work based on the previous identification of common
areas for interdisciplinary research was replicated in anthropised areas in the centre of
the island. On the basis of a list of contacts built up during previous missions and
meetings, interviews on farming practices and perceptions of nature were actually
carried out in the fields with farmers, with particular attention paid to the two central
plots of land involved in the study.

**3 Results and discussion**

**3.1 A shared history and multiple relationships**

**3.1.1 Archaeological sources**

18

The brown makis did not just arrive in Mayotte alone. Mayotte is a land of volcanic origin that emerged 8 million years ago whereas the migration of lemurs from the African continent to Madagascar (the only island they survived on) took place over 60 million years ago, probably via floating rafts that left the coast of Mozambique (Kappeler, 2000; Horvath et al., 2008). The first traces of lemurs in Mayotte date back to the ninth century AD (850-880 AD, Liszkowski, 1997) and were discovered during archaeological excavations at Dembeni. They were in the form of three half-mandibles and testify to these animals having been eaten by human populations originating from the Madagascan coast (Allibert et al., 1989). The most likely hypothesis is that these lemurs were taken as a food resource on Malagasy migrations from the north-west coast of the Grande Île. Apart from these archaeological traces there is no further information on the presence of lemurs in Mayotte over the next ten centuries. It is possible that the presence of these first traces of lemurs in Dembeni is not the sign of a durable introduction of the species and also that other makis were in fact subsequently imported.
<figure><img src="image_4.png" /><figcaption>Figure 2</figcaption></figure>

Mapping of ethnological and ecoethological study sites and brown maki countings (1999-2022).

3.1.2 Written sources

19 The travel account of an English trader named John Studdy Leigh in the first half of the nineteenth century is the first known written information regarding the presence of lemurs in Mayotte (Allibert, 1999). During Leigh's four-year expedition in the western Indian Ocean, he took great care to record the daily life of the people, the landscapes, the flora and fauna. He spent time on the island of Mayotte and observed groups of lemurs on the wooded heights during his ascent of Mount Choungi (593 m) in September 1838 (Harpet, 2011): "*They leapt from tree to tree around us, uttering their peculiar growl*" (Allibert & Verin, 1991). Leigh points out that these animals were not frightened by the presence of humans but does not mention any relations between the groups of lemurs he observed and the local populations, neither for hunting or eating purposes. Could the lemurs' real curiosity about humans as mentioned by Leigh be a sign that the human populations present at the time were not aggressive towards them? One may also speculate that contacts between human societies and lemurs were limited by the combination of dense forest cover and a human population concentrated on the coast (Gevrey, 1870-72; Allibert, 1984) and mainly involved in fishing.

20 Conversely, thirty years later in 1868 when Mayotte had been under French protectorate since 1841, François P.L. Pollen, a Dutch naturalist from the Royal Netherlands Museum of Natural History, detailed in his travel diaries (i) the French colonists' practice of hunting the 'Lemur mayottensis' with dogs, and (ii) that the animals' "flesh is excellent and tastes like rabbit meat" (Pollen & Van Dam, 1868).
Nonetheless, no information is available on the links between lemurs and the local populations of Mayotte.

### 3.1.3 Oral sources

21 From that period onwards oral sources take over. The presence of lemurs in the first half of the 20th century is mentioned in the interviews we carried out in 2000 with people in their eighties who were in their twenties when Mayotte was still a French colony in the 1940s (Harpet, 2011). These interviews revealed that lemurs were hunted in colonial times in the same way as rats, bats or wild pigs. The male hunters from Mayotte that the French colonial administration enlisted for this specific activity were paid on the basis of the number of maki tails they brought back (Harpet, 2013) but this regulated practice vanished with the successive changes in Mayotte's administrative status from 1946 onwards. This practice was part of the French hunting law system exported to the colonies during the 19th and 20th centuries. French hunting law defined wild animals as "things without masters" classified as "game" or "harmful" and considered them a source of potential physical or material damage to agricultural and forestry production (Gaidet & Aubert, 2019). This meant that preventive regulation of maki numbers was integrated into the protection and maintenance of public order (Malafosse, 1979).

22 Another local practice was still widespread all over the island at the end of the 20th century and involved young Mahoran boys catching baby makis to keep as pets. "Because makis look like humans, they're considered to be biby natanana (animals that are considered part of a house or village)" (interview, Coconi 2000, male, aged 25-30); "People enjoyed having a maki around. You can put it on your shoulder like that, it's great" (interview, Mtsamboru, 2001, male, aged 25-30). One Mahoran boy built a temporary one-room hut - a *anga* - on land belonging to his family, to mark his passage into adulthood when he left his mother's house. In this period, he used to go into the forest to catch young lemurs that had recently been weaned, using a slingshot to separate the young animal from its mother despite the clear risk of injuring or killing the targeted animals. This practice gradually disappeared at the same time as the *anga* tradition (Cassagnaud, 2006) and when the Washington Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) was implemented and which classifies the Mayotte brown maki as a protected species thus prohibiting capture, hunting and trapping.

23 In addition to these two radically different practices – the first of hunting within the framework of the colonists' exogenous regulations, and the second of domestication stemming from local culture – two mythological stories collected in 2000 (Coconi interview, 2000) describe the transformation of humans into lemurs by divine punishment because they transgressed dietary rules (using rice water to wash; misusing a ladle to serve rice). It is difficult to date the appearance of these two stories in Mayotte but we do know they are told in Madagascar and concern other species of *Eulemur* found on the main island's north-west coast in a geographical area corresponding exactly to sites where the Malagasy populations arrived in Mayotte. The data gathered during the interviews indeed give a Malagasy origin to these stories which are linked to the cult of ancestors. A dietary ban (*fady*) on eating lemurs was also linked to these two myths (*hadisi*).
24

The transformation of humans into lemurs in Mayotte was also reported in Sura VII al-Àrâf of the Koran (inserts 2 and 3) and recounted to us and translated simultaneously in 2001 (insert 4). It is very probable that this sura was adapted to Mayotte's specific context (Harpet, 2005). In the book of the Koran, only the monkey (qird) is mentioned but in the orally recited Sura the lemur (komba) appears along with the monkey so our storyteller thus evoked two distinct transformations by divine punishment - the first into a monkey and the second into a lemur, with this interpretation of the Koran shared by Koranic masters from north to south. However, it remains the prerogative of scholars.

Insert 3

<figure><img src="image_5.png" /></figure>

Sura VII al-Àrâf, verses 162-165. The mention of 'monkeys', qirada, sing. qird, is found in verse 165.
Insert 4

v. 162 : Et interroge-les sur la Cité qui était riveraine de la mer.

Quand ils célébraient le sabbat, alors les poissons [nécessaires à leur nourriture] venaient à eux, en leur jour de sabbat, sans qu'on les pêchât, alors que le jour où ils ne faisaient point sabbat, [ces poissons] ne venaient point à eux. Alors nous les éprouvions pour prix de ce qu'ils étaient pervers.

v. 163 : Et [rappelle] quand une communauté, parmi eux, demanda [à des serviteurs de Dieu] : "Pourquoi exhortez-vous un peuple qu'Allah va faire périr ou va tourmenter avec rigueur ? " [Ces serviteurs] répondirent : "Excuse à votre Seigneur ! [Nous le faisons espérant que] peut-être ils seront pieux."

v. 164 : Quand [les Infidèles] eurent oublié ce qui leur avait été rappelé, Nous sauvâmes ceux qui leur avaient interdit le mal et frappâmes d'un tourment funeste, en prix de ce qu'ils avaient été pervers, ceux qui avaient été injustes.

v. 165 : Quand [en effet] ils eurent désobéi à ce qui leur avait été interdit, Nous leur dîmes : "Soyez des singes (qirâda) abjects !²

Translation of Sura VII al-Àrâf, verses 162-165, (Blachère, 2002).

Insert 5

« Autrefois, dans un village en bord de mer, un village comme celui de M'ronabeja, vivait un peuple qui se nourrissait de sa pêche. Mais au lieu de ne pêcher que le nécessaire pour nourrir leur famille, les hommes pillaient l'Océan, tentant chaque fois de réaliser un nouvel exploit en ramenant toujours plus de poissons. En voyant cela, Allah se mit en colère et leur dit : « à partir d'aujourd'hui, vous pourrez pêcher tous les jours, sauf le samedi, jour de la chasse et de la pêche » (jour durant lequel précisément la pêche est la plus fructueuse).

En entendant ces paroles, tous les villageois ne réagirent pas de la même manière et trois groupes se formèrent.

- Le premier groupe décida de passer outre l'interdiction et continua de pêcher le samedi.

- Le deuxième groupe ne prit pas parti et laissa faire le groupe des réfractaires.

- Le troisième groupe, seul, respecta la décision de Allah.

Alors Allah punit et changea le premier groupe en singe, le deuxième groupe en lémurien. Seul le troisième groupe demeura homme. »

Interpretation of Sura VII al-Àrâf, verses 162-165, collected in August 2001.
### 3.1.4 Regulatory sources

Repeated demographic measurements of the brown maki population have been carried out in Mayotte since 1974. These studies coincided with a change in the conservation status of the species *Eulemur fulvus*, which is also the case for the original populations in north-west Madagascar. The first census in 1974 estimated the number of brown makis in Mayotte at around 50,000 individuals (Tattersall, 1977) and since then, other estimates have followed at irregular intervals, reporting significant changes to the demography and group size. Fifty years later in 2024, there were fewer than 30,000 individuals (Tarnaud, 2024). The brown maki is protected by various international and national regulations with the most important being its listing in the CITES since 1978. European Regulation 1320/2014 on the protection of wild flora and fauna by regulating trade therein also applies to this species. At the national level, Article L.411-1 of the French Environment Code and, at the departmental level, Prefectoral Order n°361/ DEAL/SEPR/2018 list the applicable protection measures with some concerning this species (Rouillé & Tarnaud, 2024).

## 3.2 The agro-ecological context: an overheated region!

### 3.2.1 From the decline in forest cover...

The topography of some sites has encouraged the preservation of original natural forests but these remain very limited (Figure 3). The woodland formations are essentially diversified and fragmented secondary forests. They are often post-agricultural forest re-growth marked by practices resulting from the transmission of family plots, which may or may not include Mahoran garden-type agroforests. From 1949 onwards, agricultural activity declined with the gradual end of rice, sisal and sugar cane production. The last sugar cane factory was closed in 1955 which coincided with a peak in agricultural decline. This led to forest regrowth, with the transfer of ownership to former workers who then developed agroforestry gardens on the former plantations. The result was that forest cover reached 65% of the island in 1987 (IUCN France, 2013).

Agroforestry gardens are often small (less than 1 ha) and difficult to distinguish from secondary forests, themselves sometimes derived from former agroforestry plots that have been abandoned but are rich in fruit species and colonised by cinnamon trees (*Cinnamomum verum* J. Presl, an invasive species). These areas are very popular with makis who can find local species and fruit trees there, including mangos. These secondary forests resulting from clearing, fire, former cultivation, agricultural abandonment and forest reconstitution make up a large proportion of Mayotte's forests nowadays. In these conditions, it is very difficult to apply the definition of woodland and this explains the variable figures because of differences in the studies and techniques used. Moreover, many wooded areas are used for agricultural practices and these may or may not be dominant. For the sake of brevity, we will focus herein on work that corresponds to the FAO definition of a forest (IGN, 2018) namely land with 10% tree cover, trees at least 5 metres tall at maturity, an area of over 0.5 hectares and an average width of at least 20 metres, and not including land that is predominantly used for agricultural or urban purposes. Alongside forests (14,100 ha), other wooded
land (3400 ha) or land with tree cover (11,800 ha) occupies as much land as actual forests (Table 2).

<figure><img src="image_6.png" /><figcaption>Figure 3</figcaption></figure>

Tree cover density. Source: Conseil général de Mayotte, DARTM, Service Ressources Forestières 2012 (based on Lidar IGN 2008 data). Areas outlined in red are owned by the State and departmental forests ([ONF, OCCM Mayotte file 20/06/2017, in Amm et al. in press].
<table><caption>Table 2</caption><thead><tr><th rowspan="2">Land use</th><th colspan="4">Area (in thousands of hectares)</th></tr><tr><th>1990</th><th>2000</th><th>2010</th><th>2015</th></tr></thead><tbody><tr><td>Forests (including mangroves and plantations) including:</td><td>19.1</td><td>15.7</td><td>14.4</td><td>14.1</td></tr><tr><td>Mangroves</td><td>0.7</td><td>0.7</td><td>0.7</td><td>0.7</td></tr><tr><td>Natural forests</td><td>-</td><td>-</td><td>-</td><td>1.1</td></tr><tr><td>Planted forests</td><td>0.2</td><td>0.3</td><td>0.5</td><td>0.5</td></tr><tr><td>Public forests</td><td>-</td><td>-</td><td>-</td><td>9.8</td></tr><tr><td>Private forests</td><td>-</td><td>-</td><td>-</td><td>4.3</td></tr><tr><td>Other wooded land</td><td>0.7</td><td>0.7</td><td>1.7</td><td>3.4</td></tr><tr><td>Total wooded areas</td><td>19.8</td><td>16.4</td><td>16.1</td><td>17.5</td></tr><tr><td>Other land including with tree cover</td><td>17.2</td><td>20.6</td><td>20.9</td><td>19.5</td></tr><tr><td>Total land area</td><td>37</td><td>37</td><td>37</td><td>37</td></tr><tr><td>Inland water</td><td>0.6</td><td>0.6</td><td>0.6</td><td>0.6</td></tr><tr><td>TOTAL</td><td>37.6</td><td>37.6</td><td>37.6</td><td>37.6</td></tr></tbody></table>

Area of forest, woodland and other land in Mayotte in 2016 according to FAO categories (IGN, 2018).

Sources: Lainé et al. 2010; AMM et al. in press.

28

Following the FAO definition, there has been a decline in forest cover since 1990 driven by anthropogenic agricultural and urban pressures (AMM et al. in press; Figure 3 and Table 2). The most recent report from the DEAL (Department of the Environment, Planning and Housing) in 2020 shows that 40% of the surface area was covered by 'natural' areas (understood here as forest reserves and anthropogenic forests with low agricultural impact) in 2016 with around 51% of agricultural areas. Between 2011 and 2016 nearly 6% of the natural areas became agricultural zones, corresponding to a reduction of over 4%. Here again though, the dividing line between forests and other wooded land is somewhat blurred in Mayotte because of the extent of areas that are close to the FAO's thresholds for tree cover and height of trees.
<figure><img src="image_7.png" /><figcaption>Figure 4</figcaption></figure>

Change in forest cover in Mayotte between 1980 and 2020. Source: FAO, 2021

29 Analysis of the density of forest cover (Figure 4) based on SPOT satellite images and aerial photographs taken regularly between 1949 and 2002 shows that 20-30% of the forest area disappeared during this period. This was also the case between 1987 and 2002 for half of the forest area because of anthropogenic pressures linked to urbanisation and accelerated clearing for agricultural purposes (DAAF, 2012; IUCN France, 2013). This degradation is tending to slow down through the development of shrub areas but nonetheless Mayotte's highly diverse and fragmented forest remains under a great deal of pressure from agriculture.

### 3.2.2 ...to the vulnerability of Mayotte's makis

30 The change in forest cover has been accompanied by profound demographic and social changes for a mammal like the maki with the average group size falling from 9 to 6 individuals since 1999. It has been estimated that the brown maki population fell by around a third between 1999 and 2020 (42,000 vs 28,500 individuals, 2020 projection based on 2016 data). Two thirds of these individuals currently live outside the island's state and departmental forest reserves in now heavily anthropised wooded areas. This population spread forms a complex mosaic with cultivated plots, particularly in the centre and north-west of the island. The density of groups of brown makis per hectare is still higher in preserved forest areas (those with a canopy at least 75% closed) but between 1999 and 2008 this increased 5-fold in cultivated areas - from 0.4 to 2 individuals per ha (Tonnabel et al., 2011; Tarnaud, 2024; Tarnaud, unpublished). This latter figure was also found in 2019 which suggests a possible stabilisation in the density of brown makis in cultivated areas during this period which can no doubt be linked to deforestation slowing in the 2010s. Whilst the population of brown makis on the island of Mayotte has drastically declined, the current number of individuals is still high in relation to the surface area occupied. This can in part be explained by the fact that groups of brown makis have extremely small home ranges of around 1 to 2 ha (Tarnaud, 2024). This was indeed observed in the two plots studied in 2021-2022 and in
several sites in Madagascar, including in natural or little-anthropised areas (Simmen et al., 2024). The makis' recognised capacity to adapt rapidly can be seen here in their diet with a few plants making up the majority of the food consumed which is made possible by the spread of food availability over time in the agroforestry context.

During the initial interviews with Mahoran farmers carried out between 2000 and 2002, the notion of the maki being an endangered species as defined by international conservation organisations was rather difficult for local people to grasp and therefore tricky to explain because the brown maki was seen as an animal they regularly encountered. In 2000, the children at Mronabeja primary school were asked to "draw the maki in its environment" and mainly chose to draw them in the agroforests (Harpet, 2002). The animal was drawn equally as much on a banana tree (antsasaka), a mango tree or on the houbouhoubou vine. The latter is commonly called "the maki fruit" while its vernacular name corresponds to two distinct species, Saba comorensis (Bojer ex A. DC. Pichon) and Ancylobotris petersiana (Klotzsch) Pierre (Figure 5). The younger generations can thus be seen to consider the maki to be a part of the fields their elders cultivate. As its presence is customary, why then should it be considered to be 'endangered'?

Twenty years on, the residents' perception of a proliferating animal is even more pronounced because of two simultaneous factors:

- The first is that the forest cover has been reduced and fragmented as human occupation and activities have steadily increased. The data show a significant loss of forest cover between 1990 and 2010, with an annual loss of almost 2%. The loss amounts to 12,000 ha since 1987 and 4700 ha between 1990 and 2010, including 3100 ha between 1990 and 2000. Meanwhile, other wooded land, including agroforests, only increased by 1000 ha between 1990 and 2010 (IUCN France, 2013). Overall this represents a significant reduction in the makis' preferred habitat, with cultivated areas now replacing it.

- Secondly, the reduction in natural forest areas have made makis more likely to forage in wooded areas made up of agroforests or former fruit tree plantations, where food is abundant. This has meant the maki is more visible so that, while the overall impression one may have is that the number of makis is increasing, in fact the censuses show a steady decline.
<figure><img src="image_8.png" /><figcaption>Figure 5</figcaption></figure>

Child's crayon drawing of makis on wild and cultivated fruit trees, 2001, Mronabeja School.

### 3.2.3 ...through changes in human representations and behaviour

33 Human societies have developed multiple relationships with the brown makis of Mayotte (hunting, consumption, domestication, protection) and correspond to specific eras and cultures. The last relationship cited, that of "protection", was mainly asserted at the end of the twentieth century by associative networks mainly represented by the metropolitan population of Mayotte. This relationship sometimes brought two cultures into virulent opposition. These are: 1/ the native population's so-called local culture that is attached to using religious and symbolic precepts to define its relationship with living creatures, and 2/ the 'Mzungu' (metropolitan, ex-colonial) culture supported by a new State with its rule of law promoting the protection of biodiversity. Indeed, the latter philosophy led to the Terre d'Asile association transporting around a hundred brown makis that had been found injured or trafficked to the Mbouzi islet between 1997 and 2004. This islet was classified as a National Nature Reserve (RNN) in 2007 (the first in Mayotte) which meant the association had to cease its activities. In 2023, the population of brown makis on the islet of Mbouzi was estimated at just under 200 individuals and was dynamic, with annual births observed (Tarnaud, 2023). The reserve's island configuration undoubtedly makes the preservation of makis easier and this is also the case as regards the maintenance of a minimally anthropised ecosystem easier.

34 Conversely, at the time of Grande Terre's 'departmentalisation' in 2010, conflicts between farmers and brown makis in their fields on the island were reported in interviews. Makis were, for example, described as "wasters" of fruit crops and "worse" than rats which "at least eat the whole fruit before starting to eat another one" (Interview,
Coconi, October 27th 2010). The maki is not the only 'pilferer' in the fields but it is still the toughest to the extent that local farmers noticed its diet had changed, reporting that "it eats green bananas" (Interview, Mayotte, Ongojou, October 26th 2010). The once-popular strategy of cutting still-green banana bunches so these could ripen away from thieving makis is no longer the norm. The 5th census in 2010 of Mayotte's brown makis confirmed the decline in this species but the information gathered from farmers starkly contrasted with the figures announced with farmers reporting that "there are more makis" or "we see a lot more...". (Mayotte interview, Coconi, October 20th 2010; Hajangoua, October 21st 2010). In reality, these results are only antagonistic in appearance because makis actually are more numerous in the fields. However in the course of our discussions, the farmers readily acknowledged that numbers of makis are declining in the forest because "nothing grows in the forest anymore, they've cut everything down" while observing that "before, when there was more forest, they didn't come here". This means that lemurs therefore have to "come down to feed" (Mayotte interview, Mtsangamoudji, October 27th 2010).

In 2000, when the farmers were asked "how do you keep the makis away when they steal from your field?" most replied that "you mustn't bother them because their fate is bad enough" (Coconi interview, July 2000) which is a reference to mythological tales. Scarecrows were sometimes made from sheets and if a group of makis was spotted in the agroforests, children took it upon themselves to scare them away from the crops, knowing full well they would be back the next day. By 2010, the threshold of tolerance had changed, with many of the respondents reporting that the makis' behaviour had also changed. "The makis have changed as well", "they come into houses and are aggressive sometimes" (Mayotte interview, Mronabeja, October 31st 2010). The anecdotes of lemur attacks respondents related were found to include much harsher judgements when children are involved.

In 2002, the farmers still considered the presence of makis in the cultivated areas to be beneficial. In fact the maki played a kind of substitute role when humans were not present within these 'semi-cultivated' or 'semi-spontaneous' areas but this consideration subsequently became increasingly more marginal. In 2024 the maki was still accorded a 'special' status in local representations in Mayotte, namely that of a human transformed by divine punishment into a maki who must be spared and allowed to share the benefits of abundant nature. The local population, particularly the younger educated generations who are influenced by Western lifestyles, are moving away from such ancestral precepts and beliefs so that these beliefs are no longer adapted to a market economy, despite still prevailing at the end of the 20th century. Traditional subsistence farming is being replaced by larger-scale cash crop farming of products with greater export value. Local people who used to be attentive observers are now becoming active farmers. This change of model has had direct repercussions on how agroforestry space is cultivated and conceived of.

At the very start of the 2000s, nearly two-thirds of the island's households were involved in agriculture on 20,250 ha of land (Sourisseau et al., 2008). In 2010, the recorded number of farming households was close to the previous figure (15,700 farming households) (DAAF, 2012). In 2020, the departmentalisation process led to the census criteria being changed to comply with the CAP (Common Agricultural Policy) model. The term 'agricultural household' was replaced by 'agricultural holding' to bring this aspect into line with the census protocol used in France's 101 departments.
The latest Agricultural Census (RA) found that there are now 4315 farms covering an area of around 6000 hectares. This apparent fall in the area cultivated during this 10-year period can more specifically be explained by the use of a different counting method to the one used for the previous census (between 2008 and 2010). The new method no longer takes agricultural plots used exclusively for food production into account and only counts farms that market all or part of their production. This census process marks a turning point in the representation of Mayotte's farming model in Mayotte and in what government departments now consider to be 'agricultural activity'.

This new approach to farming has indirectly led to wild frugivorous species like the maki now being perceived as potential 'pests' whereas they were previously tolerated or even appreciated in a subsistence society where farming was a family activity.

Also, there has been a change since 2010 to the frequency of the presence of farmers in the agroforestry gardens. While weekends in these convivial spaces remain lively and festive, we no longer, or only sporadically, find people who go to the fields early in the morning when it is still cool to spend the whole day there (participant observation, field monitoring, Msangamouji - Mronabeja, 2000 and 2010) which was the case in the early 2000s.

In 2024, two-thirds of the farmers were aged over 60. Nearly a third of the population of Mayotte is involved in farming but it is a secondary activity for the majority of working people. The new generations of farmers are adopting new habits that change their relationships with the land and thus their previously close relationship with makis.

The changes to the usages and perceptions of the maki (its status in local culture and its increasingly problematic place in anthropised areas) are not unrelated to the profound changes to the status of Mayotte's farmers and their relationship with cultivated areas. Interviews carried out in 2021-22 revealed a clear divergence between the responses of cash crop farmers who consider the maki a pest and those of strictly food crop farmers (Lecarpentier, in press). In fact, the curve of tolerance towards makis and other wild frugivorous species depends on the farming model practised.

### 3.3 The adaptation and survival strategies of humans and makis

When our initial research began in 1999, we had not realised the extent of the rapid environmental changes the territory of Mayotte would undergo because of successive geopolitical and socio-demographic transformations. Between the second half of the nineteenth century and the beginning of the twenty-first, Mayotte's institutional history was particularly turbulent with the island successively passing from the status of colony to that of an Overseas Territory (TOM) and then to that of Overseas Department (DOM) in 2011 (Béringer, 2012). This DOM status announced in the 1990s with a 10-year assimilation programme made the island of Mayotte the 101st French department and the 5th overseas department but profound structural upheavals have accompanied this process of change. In 2002, a multi-disciplinary (archaeology, history, linguistics, anthropology and psychology) study of the island's situation at the beginning of the 21st century highlighted "*changes that have been made without sufficient transition periods*" to ensure continuity and social coherence within the territory (Allibert, 2002). Twenty years later, the effects of the island's brutal political and economic changes are now glaringly apparent and include crises related to water,
governance, migration and agriculture against a more global backdrop of climate crisis. All of these crises have led to worrying repercussions on the island's natural areas and biodiversity. Natural environments are deteriorating and leading to the increase we are seeing in the number of animal species subject to strong anthropogenic pressures, particularly the maki which is already classified as a "vulnerable" species according to the IUCN criteria in 2018. How, then, are humans and makis adapting to these changes?

### 3.3.1 Humans

43

The Malagasy myths and the sura from the Koran mentioned earlier helped organise and maintain a traditional society based on direct access to and consumption of resources produced locally. These included warnings against human excesses like overusing the resources God lavishes on mankind (sura from the Koran) and wasting food (mythological accounts). These precepts undoubtedly had a greater influence on behaviour than is the case today. Their application tended to regulate access to resources in a self-subsistence society, helping to preserve the island's ecosystems. Traditional rules stipulated that it was forbidden to hunt or eat makis.

44

The latest ethnographic survey was run in 2021-2022 and confirmed trends that had already begun in 2010 - the larger the cultivated area and the less diverse the fruit species (e.g. banana monoculture), the less animal species are tolerated. Farmers implement more numerous and systematic adaptation strategies to target predators. These include replacing fruit species, planting plant barriers, and putting up wire fences several metres high around cultivated areas. They also employ security guards and even use guard dogs which is an unexpected choice in a society in which dogs are not viewed positively at all. This is the case of the Tsararano plot's owner who uses a combination of wire fencing, temporary human surveillance and the 24/7 presence of guard dogs.

45

The choice of a given control strategy is correlated with the type of farming involved and the amount of time devoted to this activity. When farming becomes the main income-generating activity, farmers tend to adopt a more defensive posture, guarding against theft, rather than their previous 'laissez-faire' attitude. On the most exposed plots, this posture can extend to active defence measures including the use of chemical products to get rid of pests (Lecarpentier, in press). Farmers whose business model entails selling all or part of their produce aim to expand and specialise and so are breaking away from the traditional 'mix' system practised on small plots of land that are ideal for growing subsistence crops. The growing intolerance towards wild species is linked to the monoculture model and intensive agriculture imported from mainland France where the same issues have been encountered. For over a century, common agricultural policies have imposed a single agricultural model on France's overseas territories. For example, 81% of the arable land on La Réunion Island is taken up by industrial crops that originated from sugar plantations (Agreste, 2022). While Mayotte currently has only 1%, this figure is rising. More sober and virtuous models like permaculture which is closer to the agroforestry model are having a trouble in finding their place. In 2020, only 22 of the 4315 farmers surveyed reported that they practiced organic farming (Agreste, 2021).
### 3.3.2 Makis

Like other phylogenetically related *Eulemur* species, *Eulemur fulvus* displays a considerable level of ecological flexibility which is probably due to its low energy requirements (Simmen *et al.*, 2010; 2024). These lemurs can adapt effectively to variations to their environment including the presence of humans. Although their preference is to live in trees, one clear sign of this adaptability is their capacity to move into more or less open agroforestry systems, resulting in a more terrestrial mode of locomotion. By 2020, it was estimated that two thirds of the population of brown makis live outside Mayotte's departmental and state forest areas in wooded areas with at least 50% of their canopies open (Tarnaud, 2024). The 2021 results showed that there were more groups of brown makis living in cultivated areas than in forests so clearly the trend has been reversed in under 10 years.

In the two agricultural plots studied in detail, brown makis are almost entirely dependent on cultivated resources. Indeed they spend over 85% of their feeding time on these resources, depending on the season and the plot of land. The resources concerned are mainly fruits, including bananas, guavas, cytherea plums, papayas, carambola, various Annonaceae, jackfruit and certain citrus fruits. The spatial concentration of resources and fruit varieties the makis select for their nutritional and organoleptic qualities means that they spend less than 30% of the day feeding and foraging to meet their metabolic requirements. This dietary dependence is actually even more problematic because if massive crop protection is implemented, the plots will not be located close enough to the forest reserves for the groups to adjust their foraging strategy effectively. Groups of makis that occupy areas at the intersection of forested zones and neighbouring agricultural zones have an ecological relationship with humans that we may call facultative commensalism if we accept the notion of limited impact for farmers who say they tolerate makis. However, even in a scenario involving relatively exclusive home ranges, such as those observed on the two control plots in 2021-2022, intense competition for food with groups of makis on the forest edge is to be expected if crop protection becomes the norm. As there are respectively 63 and 42 individuals in each of the two plots, considerable population densities are reached. Nonetheless, these are probably not typical of the majority of cultivated areas, which are generally less than 1 ha in size and with an estimated brown maki density 2.6 times lower than observed in forest areas with a 75% closed canopy. Cash crop plots are an exception to this as they are larger than plots used for food crops (in 2010, only 2% of plots covered more than 2 ha in Mayotte; Agreste, 2011). Therefore, it is to be expected that exclusive protection of these plots would negatively impact the reproduction and survival rates of many groups. This phenomenon is continually increasing and represents a profound change in human/environment relations in Mayotte and de facto in human/lemur relations. The conservation of makis is inevitably facing repercussions due to the growing perception that they are predators of fruit crops.

## 4 Conclusion

Unlike its Malagasy conspecifics, *Eulemur fulvus* in Mayotte has never lived without humans being present. The makis in Mayotte were introduced by humans from the
island of Madagascar and have been part of the landscape in Mayotte for at least twelve
centuries. The maki mainly inhabited the island's strictly forested areas, which
provided most of its food. However, there has been a sharp fall in woodland cover from
65% to 37%, which has also transformed the relationship between humans and their
environment.

49

International ecological conservation arguments are relatively inaudible in what is France's poorest department. Seventy-seven percent of the population of Mayotte lives below the national poverty line, which is five times higher than in mainland France (https://www.insee.fr/fr/statistiques/4632225). If solutions are not developed to ensure the coexistence of *Eulemur fulvus* with human societies on the island, makis will eventually disappear from shared spaces. So, what would we lose if the Mayotte brown lemur disappeared? As frugivorous animals that disperse seeds, makis have modified the forest landscape over the centuries, particularly by favouring the dispersal of cultivated plants. If makis were to disappear, other species that can play a comparable functional role in the regeneration dynamics of ecosystems like the large fruit bats that are very abundant in Mayotte could prove effective substitutes for certain zoochore plant species. However, recolonising secondary forest habitats with native plant species effectively require an assemblage of terrestrial and flying fruit-eating wildlife, in which makis play an essential role in disseminating seeds.

50

One of the radical solutions that some local people have put forward to solve the "maki problem" is to isolate the species in protected island areas like islet of Mbouzi, which has been classified as a 'National Nature Reserve' (RNN162) since 2007 - "They have their island, so why should we put up with them here?" (interview with farmer, Coconi 2015). This position is similar to that of some conservation biologists and consists of the complete separation of biodiversity protection zones and farmland which has certainly proved effective in terms of agricultural production yields (Phalan et al., 2011). However, this overlooks the importance of the ecosystemic services biodiversity provides for rural societies and also restricts nature to the role of a mere food resource (Bahuchet, 2017). In our case, it reduces the Mayotte brown maki's place and role to just its ecological and recreational function and does not take into account the maki's symbolic and structural place in the culture of the island. In traditional Mahoran society, the maki is considered to be a reference point for the maintenance of a rich, healthy and abundant biodiversity. Its presence in agroforests guarantees the good health of the woodland as gardeners well know. Indeed they encourage such interactions and this socio-ecological complicity by maintaining animal and plant diversity on their plots of land. Until recently, the Mahorans (men and women alike) defined themselves as "Muslim residents of Mayotte with a field for their subsistence" (Blanchy, 2002) and their attachment to their plots of cultivated land was a fundamental feature of Mayotte culture. In this way, the disappearance of the brown maki in Mayotte would symbolise the end of a complete agroforestry system based on a balance between nature, the supernatural, and humans, upon which the local, self-subsistence economy was built.

51

Intensive monoculture would become the norm and the role of the gardener would disappear to be replaced by that of the farmer modelled on the French mainland. The disappearance of makis would also go completely against Article 1 of the French Rural and Maritime Fishing Code dated July 1st 2023. This explicitly supports the agro-ecological model "based on biological interactions and the use of ecosystem services and the potential offered by natural resources" (https://www.legifrance.gouv.fr/codes/article_lc/
LEGIARTI000043978760). The model of intensive, homogeneous monoculture agriculture has demonstrated its highly negative effects on biodiversity and, more broadly, on the health of ecosystems and humans (depletion and pollution of soils through the use of pesticides, fragmentation of ecological habitats and so on). When compared with the environmental situation of other French overseas island departments, Mayotte, the 101st French department, has considerable ecological potential. Mayotte was a latecomer to the departmentalisation process and is definitely lagging behind in terms of economic development as defined by GDP (Roinsard, 2022). Conversely its intermediary position halfway between tradition and modernity offers unique agro-ecological prospects which other overseas departments that have already gone through the post-war agro-industrial revolution would have difficulty equalling. Twenty years ago, agricultural policymakers viewed Mayotte's difficult geography, small plot sizes, fragile soils, lack of means of production and absence of mechanisation, etc.) as weakness. However, in an era of profound questioning of the intensive agriculture model, these characteristics could be transformed into real opportunities (Huet, 2021). Today, Mayotte's younger generations are more inclined to follow the mainland model but nonetheless remain deeply attached to the family agroforestry gardens of the past. These gardens offer them a quality of life and aesthetic pleasure, as well as food resources, away from the bustling cities (Longépée et al., 2019). In an attempt to capitalise on this trend among young people in Mayotte Mayotte's Directorate of Food, Agriculture and Forestry launched a call for projects (https://www.mayotte.gouv.fr/contenu/telechargement/8859/67696/file/AAP_janvier_2018.pdf). Its aim was to (i) encourage the creation of economic and environmental interest groups (GIEE) for "the promotion of agroecology and agroforestry in Mayotte" and (ii) attract and train Mayotte's young generations in agroecological practices.

52 Only through rural development that cares for biodiversity and humanity alike, involving a triple alliance between agronomic, social and ecological sciences, can Mayotte's brown makis continue to coexist with humans.

## Acknowledgements

53 We would like to thank the François Sommer Foundation for funding the "Human-wildlife conflicts" project from 2021-2023. This project was made possible by the agreement of the farmers of the two plots we studied and by the participation in data collection and analysis of Maëva Lecarpentier, Laurane Rami, Guillaume Correa-Pimpao and Romane Ibanez. We would also like to thank the DEAL of Mayotte, the *Conservatoire botanique des Mascarins*, the ONF and the DEDDE with Mahamoud Ahmed-Omar for supporting us in this programme.

## Declaration of absence of conflict of interest

54 The authors declare they have no conflicts of interest.
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ABSTRACTS

One of Mayotte's most emblematic wildlife species is the brown lemur *Eulemur fulvus*, introduced from Madagascar by human populations during the first migrations from the 8th century onwards. Long valued and protected in local culture, it is now seen as a threat to agriculture in a
society that is changing as a result of departmentalization. Growing pressure on forest areas is forcing lemurs to forage in agroforests and orchards, thereby altering their relationship with human populations, as well as their distribution in protected forest areas and cultivated zones. Increasingly regarded as pests, their fate is more than ever dependent on human activities in a local context of socio-political, cultural and environmental crisis. This article looks back over 25 years of interdisciplinary research in the humanities, social sciences and life sciences, with an update on the ecological history of *Eulemur fulvus* and human/lemur interactions on the island of Mayotte.

L'île de Mayotte compte, parmi ses espèces faunistiques emblématiques, le maki brun *Eulemur fulvus* introduit depuis Madagascar par les populations humaines lors des premières migrations à partir du VIIIe siècle. Valorisé et protégé dans la culture locale, il est aujourd'hui considéré comme une menace pour l'agriculture au sein d'une société qui se transforme sous l'effet de la départementalisation. Une pression grandissante sur les espaces forestiers pousse les makis bruns à se nourrir dans les agroforêts, modifiant par là même leurs relations avec les populations humaines, de même que leur répartition dans les aires de forêts protégées et les zones cultivées. Leur sort est plus que jamais dépendant des activités humaines dans un contexte local de crise sociopolitique, culturelle et environnementale. Cet article propose de revenir sur 25 années de recherche interdisciplinaire en sciences humaines et sociales et en sciences du vivant, en actualisant la connaissance de l'histoire écologique d*Eulemur fulvus* et des interactions humains/lémuriens sur l'île de Mayotte.

## INDEX

**Mots-clés:** Eulemur fulvus, maki brun, déforestation, agroforêts, représentations, pratiques.

**Keywords:** Eulemur fulvus, brown lemur, deforestation, agroforests, representations, practices.

## AUTHORS

### CLAIRE HARPET

UMR 5600, Environment, City and Society Laboratory, Lyon 3 University, Lyon, France
UMR 7206, Eco-anthropology Laboratory, CNRS/MNHN/Université Paris Cité, France
Corresponding author: claire.harpet@univ-lyon3.fr

### LAURENT TARNAUD

UMR 7206, Eco-anthropology Laboratory, CNRS/MNHN/Université Paris Cité, France

### BERNARD RIÉRA

UMR 7179, Adaptive Mechanisms and Evolution Laboratory, CNRS/MNHN, Brunoy, France

### BRUNO SIMMEN

UMR 7206, Eco-anthropology Laboratory, CNRS/MNHN/Université Paris Cité, France