PhD Verohanitra M Rafidison

PhD Bakolimalala Rakouth
email: ba.rakouth@yahoo.fr

Stéphanie Carriere
email: stephanie.carriere@ird.fr

PhD Finn Kjellberg
email: finn.kjellberg@cefe.cnrs.fr

PhD Yildiz Aumeeruddy-Thomas

PhD Stéphanie M Carrière

Multiple values of isolated and clusters of

Keywords: Ficus, Madagascar, local knowledge, social-ecological systems, traditional practices, forest corridor, biodiversity, collaborative approaches +33643118153

published or not. The documents may come

approaches which consider synergies between local and scientific knowledge.

Response to Reviewers:

Responses to reviewer 1: Many thanks for your detailed comments and your thoughtful visions that have helped us clarify our statements. Please find below our responses: 1."you repeatedly talk about "noble", "poor" and "ordinary" social classes who have different access rights to different Ficus trees. I am aware that in many Malagasy societies caste systems are in place but many readers are not, so i would suggest that you explain this early in the introduction and also relate these groups ethnically, in power and its socio-cultural context".

We have responded to this comment. Please see lines 92-127.

2. "You don't really study anything about livelihoods, therefore, i would remove this from all parts of the manuscript, i.e. for sustaining "biodiversity and livelihoods", it is certainly not the case that this is always mutually beneficial.Yu would have had to give this a framework , i.e. "sustainable livelihoods" or similar but you do don't. As it is now , it is not clear what is meant. Livelihoods mean different things to economists, geographers and social scientists".

In the introduction (lines 81-94), we explain the following : (1) that livelihoods are very strongly linked in Madagascar to biodiversity. We explain that human activities in Madagascar have driven deforestation. There is thus no ambiguity in our statements.

We do not imply that biodiversity and livelihoods necessarily have positive effects on each other. When we say the following : « The current decline in the rule of law in Madagascar could greatly harm biodiversity and local livelihoods", we clearly mean that local livelihoods are highly dependent upon biodiversity. We do not say that local livelihoods necessarily sustain biodiversity. The whole paper in fact aims at identifying specific connections between people and some elements of biodiversity, in this case Ficus species, that could potentially help sustain biodiversity while also sustaining livelihoods. The roles of Ficus as keystone species that can potentially enhance the conservation of biodiversity are fully recognized in literature that we mention.

Furthermore, as ethnoecologists, and not "social scientists" ( ethnoecology combines ethnology and approaches in ecology), we do not consider that livelihoods are independent from ways people perceive, use, and manage Ficus species within their rural landscapes. In this particular case, these elements of biodiversity have impacts on soil (fertilization) and water, impacts on feeding habits of children (much work on food systems shows that children access foods that adults do not use and that this often neglected part of their diets contributes nutrients essential to their health), on culture and interpersonal relationships, and on people's beliefs and religion. It is fully recognized that human well-being is linked to all these dimensions of livelihood (cf Sustainable Development Goals, also see Sterling et al. 2017). We therefore consider that our use of the term "livelihood" fits with the content of this paper and cannot be removed.

Sterling, E. J., Filardi, C., Toomey, A., Sigouin, A., Betley, E., Gazit, N., … Stege, K. (2017). sustainability indicators across scales. Nature Ecology & Evolution, 1(December). https://doi.org/10.1038/s41559-017-0349-6

3. « the frameworks about "nature" and "biodiversity" are inconsistently used throughout the manuscript. Given the (partly) social science approach of this paper, this needs more clarity and I would expect this section to be a bit more critical. Nature is a social construct, according to most social scientists, whereas biodiversity is not, which is much narrower defined but also includes many other things beyond tree diversity »As ethnoecologists, we do not use « Nature » only as a social construct, but we use a precise definition (given on lines 96 -100) and we explain how the concept of nature that we use relates to biodiversity. We do not wish to remove the word biodiversity from the paper, because it is clear that when we speak of a species of Ficus with a scientific name, its position in a landscape, and how people relate to it, we do consider it as an element of biodiversity to which people refer to very precisely, even though vernacular classifications may differ from scientific ones.

Malagasy rituals, traditions and beliefs are around nature but NOT about biodiversity, this is an interpretation, and many anthropologists and social scientists will not agree with this statements. Either remove it or back it up with more discussion and references. Explained above and references added in the manuscript.

INTRODUCTION

Madagascar is widely recognized for the high rate of endemism and the richness of its biodiversity [START_REF] Goodman | The natural history of Madagascar[END_REF][START_REF] Ganzhorn | Explaining Madagascar's biodiversity[END_REF]). Most of the population of Madagascar (80 %) lives in rural areas, conducting agriculture and pastoralism, and relies on biodiversity for food, medicine, fuelwood, and timber [START_REF] Waeber | Dry forests in Madagascar : neglected and under pressure[END_REF]. Increasing threats to Madagascar's biodiversity critically affect the integrity of ecosystems, inside protected areas and in rural landscapes, affecting local livelihoods [START_REF] Jones | Last chance for Madagascar's biodiversity. Nature Sustainability[END_REF]. Agriculture, charcoal production and zebu grazing in forest areas by local inhabitants have driven deforestation, resulting in loss of habitats and biodiversity [START_REF] Waeber | Dry forests in Madagascar : neglected and under pressure[END_REF]. However, corruption associated with new agribusiness and mining activities brings new threats from forces acting at national and global scales. The current decline in the rule of law in Madagascar could greatly harm biodiversity and local livelihoods [START_REF] Jones | Last chance for Madagascar's biodiversity. Nature Sustainability[END_REF].

In this paper, nature is used following the concept of Nature's Contributions to People (NCP) defined by Diaz et al (2018) whereby nature relates to living nature, i.e., the diversity of organisms and the ecological and evolutionary processes within ecosystems. "Nature" can have different cultural meanings for different societies. In this paper, we consider this living part of nature as equivalent to biodiversity.

In Madagascar, people are connected to the power of life and nature known as Hasina, through the mediation of ancestors, their burial places and particular species or elements of ecosystems [START_REF] Harper | Memories of ancestry in the forests of Madagascar[END_REF], Aumeeruddy-Thomas et al. 2018). Access to different parts of territories and resources follows specific rules and management practices framed upon reciprocity between people and elements of biodiversity [START_REF] Diver | Recognizing "reciprocal relations" to restore community access to land and water[END_REF]. Within this context, some species, groups of species, or ecosystems such as portions of forests are seen as active agents that may interact with people or elements that participate in the identity of people. Indeed, from the perspective of some Malagasy groups, forests reflect not only natural history, but social history. They are viewed as symbols of shared histories of collective empowerment, including resistance to outside control [START_REF] Harper | Memories of ancestry in the forests of Madagascar[END_REF]. Social organization governing access to different types of elements of nature in Madagascar is based on social hierarchies and systems of power that have built up along history. In particular, the Merina 1 king Andrianampoinimerina ((1787-1810) , united Malagasy groups (e.g. Sakalava, Betsileo, Betsimisaraka and many others that previously were organized in the form of small independant kingdoms) to establish a state at the beginning of the 19 th century. enforced a caste system including andriana (noble), hova (free) or andevo (slave). This system served the interests of the king's taxation system based on a reinforced link to the land and tombs that also facilitated the control of allegiance by all social groups to the Merina. The andriana status could be conferred for loyalty or service to the king, while hova took on varied meanings in different parts of the island but in all cases referred to people possessing lands and ancestors buried in these lands.

Some Ficus species, such as Ficus lutea, are among these particular species that symbolized the power of the king Andrianampoinimerina and contributed to enforce his hegemony (Aumeeruddy-Thomas et al. 2018).

In the Highands, the organization of landscapes is defined by diverse categories of spaces, which each bear a local term that designates jointly ecological aspects and associated As in the rest of the world, since the 1970's there have been many efforts in Madagascar to develop approaches that integrate biological conservation and human development [START_REF] Marie | Taking into account local practices and indigenous knowledge in an emergency conservation context in Madagascar[END_REF][START_REF] Aumeeruddy-Thomas | Savoirs locaux et biodiversité : interactions sociétés et aires protégées. Etudes comparatives : Indonésie, Népal, Pakistan et France[END_REF]. Numerous efforts were made in Madagascar by a large diversity of NGOs to integrate human development into conservation. However, it seems that the urgency for developing the conservation network, led by the global conservation agenda and its priorities, accorded little attention to local knowledge, practices and institutions including attachment to land [START_REF] Marie | Taking into account local practices and indigenous knowledge in an emergency conservation context in Madagascar[END_REF]. Globally, results of similar efforts elsewhere have generally been unsatisfactory [START_REF] Ward | Protected area co-management and perceived livelihood impacts[END_REF]. In Madagascar, GELOSE, a national law adopted in 1996, aimed to devolve the management and conservation of forests to local communities and stimulate economic development. Contractual agreements between the state and local community-based groups (CoBa) were drawn up, defining rules of access in a zonation system based mainly on conservation priorities. Social scientists pointed out that GELOSE redefined access to spaces in a way that did not take into account practices of the area's rural population [START_REF] Blanc-Pamard | L'illusion participative, exemples ouest-malgaches[END_REF]. [START_REF] Mcconnell | Challenges of forest governance in Madagascar[END_REF] further explain: The policies "reify monolithic communities to which authority can be rapidly devolved, when in fact these 'local communities' are difficult to define, much less successfully engage in conservation activities, especially after a long era of strong state control". These experiences highlight the gap between global conservation endeavors and local socialecological dynamics.

At the global level, new approaches take into account the importance and role of biodiversity outside protected areas, and address landscape and social dynamics [START_REF] Mora | Ongoing global biodiversity loss and the need to move beyond protected areas: a review of the technical and practical shortcomings of protected areas on land and sea[END_REF]. New types of conservation areas include IUCN categories V and VI, UNESCO Maintaining extensive anthropic landscapes featuring a mosaic of low-input agriculture has been characterized as a "land-sharing" strategy that conserves biodiversity while sustaining agricultural production [START_REF] Perfecto | The agroecological matrix as alternative to the landsparing / agriculture intensification model[END_REF]. Such landscapes are often significant reservoirs of biodiversity [START_REF] Kun | Intermediate landscape disturbance maximizes metapopulation density[END_REF][START_REF] Kjellberg | Corridor de végétation et conservation d'un groupe clé de voute de la biodiversité, le cas des Ficus et des communautés associées[END_REF]. This biodiversity may be critical to ensure the resilience of ecosystem functions, particularly in the present context of rapid climate change [START_REF] Manning | Transferring biodiversity-ecosystem function research to the management of 'real-world' ecosystems[END_REF][START_REF] Renard | National food production stabilized by crop diversity[END_REF].

Tees or clusters of trees (e.g. small woods, hedges) initially part of forest environments, or those planted as isolated trees or within clusters of trees, can help create or re-establish connectivity in fragmented a agrarian landscapes. They offer resources to animals and accelerate plant successions [START_REF] Carrière | Orphan trees of the forest. Why do Ntumu farmers of southern Cameroon protect trees in their swidden fields[END_REF][START_REF] Manning | Scattered trees are keystone structures: Implications for conservation[END_REF][START_REF] Eden | Isolated Ficus trees deliver dual conservation and development benefits in a rural landscape[END_REF]. Species of Ficus (Moraceae) are particularly important in this regard. Trees of this pantropical genus, which numbers about 800 species, are hubs in vast webs of interactions. These include the specialized insect pollinators, parasites and parasitoids of Ficus. They also include mammals and birds that eat fruits and disperse seeds not only of Ficus but also many other plants [START_REF] Shanahan | Fig-eating by vertebrate frugivores: a global review[END_REF][START_REF] Harrison | Seventy-seven ways to be a fig: an overview of adiverse assemblage offigs in Borneo[END_REF][START_REF] Kjellberg | Biology, ecology and evolution of fig-pollinating wasps (Chalcidoidea, Agaonidae)[END_REF]. When Ficus grow as isolated trees, they act as stepping-stones for forest animals in open habitats [START_REF] Martin | Conservation value for birds of traditionally managed isolated trees in an agricultural landscape of Madagascar[END_REF]. They also function as nuclei accelerating plant succession, as the frugivorous animals that visit them disperse seeds of forest plants in their droppings. These seeds germinate and establish, forming small patches of vegetation around these focal trees [START_REF] Carrière | Orphan trees of the forest. Why do Ntumu farmers of southern Cameroon protect trees in their swidden fields[END_REF]. In Madagascar and elsewhere, Ficus trees are also associated with multiple social values that connect people to territories and nature (William and William 2013, [START_REF] Eden | Isolated Ficus trees deliver dual conservation and development benefits in a rural landscape[END_REF][START_REF] Rafidison | Pratiques et connaissances naturalistes des communautés Betsileo: lisière du corridor forestier Andringitra-Ranomafana, Madagascar[END_REF][START_REF] Aumeeruddy-Thomas | Ongoing domestication and diversification in grafted olive-oleaster agroecosystems in Northern Morocco[END_REF], 2018). Madagascar has 25 native Ficus species belonging to four subgenera, of which 15 species are endemic and four are shared with the African continent [START_REF] Dalecky | Moraceae, Ficus and associated fauna[END_REF][START_REF] Rafidison | Ethnobiologie et écologie des Ficus des terroirs Betsileo et du corridor Ranomafana -Andringitra (Madagascar). Dissertation University Antananarivo Rafidison[END_REF]. Taxonomic and ecological information on Ficus species of Madagascar is given in Table 1. Three Ficus species represent 33% of the isolated trees (sample size 45) in Betsileo rural landscapes; they attract numerous bird species as well as bats [START_REF] Martin | Conservation value for birds of traditionally managed isolated trees in an agricultural landscape of Madagascar[END_REF]. Some species, such as F. tiliifolia, are highly attractive to frugivorous animals [START_REF] Goodman | Rarity of figs (Ficus) on Madagascar and its relationship to a depauperate frugivore community[END_REF]. Ficus have a cultural value for the Betsileo, and are protected for this reason, but a full understanding of their multiple values and their contributions to social-ecological dynamics is still lacking [START_REF] Moreau | Les gens de la lisière[END_REF][START_REF] Martin | Conservation value for birds of traditionally managed isolated trees in an agricultural landscape of Madagascar[END_REF].

OBJECTIVE

Our objective is to understand social-ecological dynamics that affect the distribution, uses and values at the landscape level of Ficus species that are protected by Betsileo communities in two rural landscapes. We use our findings to discuss the importance of collaborative approaches that consider local social-ecological dynamics that can improve biodiversity management and human livelihoods.

We focus on farmers' local knowledge and historical, cultural and practical reasons that explain why farmers plant and/or protect Ficus trees in their agrarian landscape. We use an ethnobotanical approach to assess the uses, multiple values and distribution of Ficus spp. within the territories of two Betsileo villages that border the Ambositra-Vondrozo forested corridor.

Our specific aims are to:

(1) Identify the relationships between their distributions and uses, within the history of the construction of their rural landscape by the Betsileo in two sites;

(2) Identify past and present socio-political, economic, practical and symbolic values attributed to Ficus species by two Betsileo communities and consequences of these values for their management;

(3) Propose rules of thumb for designing collaborative approaches based on our understanding of local social-ecological systems and the roles of key elements such as Ficus species.

STUDY SITE, MATERIALS AND METHODS

Our study area is located on the eastern side of the Malagasy Highlands. It is bounded to the west by a mountain ridge that stretches north to south along Madagascar. A forest strip known as the Ambositra-Vondrozo corridor, characterized by a dense mid-altitude humid forest (Goodman & Razafidratsita 2001) et al 2010[START_REF] Rafidison | Ethnobiologie et écologie des Ficus des terroirs Betsileo et du corridor Ranomafana -Andringitra (Madagascar). Dissertation University Antananarivo Rafidison[END_REF]. Choice of these two sites for this study allowed investigating how social values accorded to Ficus species interact with their ecology.

Our two study sites belonged historically to distinct Betsileo chiefdoms until the unification of the country by the Merina in the 19 th century. The Betsileo cultural group the largest cultural group that inhabit the southern part of the Highlands [START_REF] Harper | Memories of ancestry in the forests of Madagascar[END_REF]. The Betsileo originate from an admixture of groups of migrants who successively colonized the highlands. Their language includes linguistic elements of different cultural groups (Antemorro, Tanala, Merina) [START_REF] Dubois | Monographie des Betsileos[END_REF]. They transformed the forest with swidden agriculture (tavy), cultivating mostly rainfed rice [START_REF] Dubois | Monographie des Betsileos[END_REF]. The advent of irrigated rice cultivation among the Betsileo during the late 17 th and early 18th centuries has been attributed to internal changes associated with a demographic increase, participation in trade of slaves and weapons, and contacts with the Merina [START_REF] Kottak | The process of state formation in Madagascar[END_REF]. The Betsileo were comprised of distinct small chiefdoms, each with its specific frontiers, territories and hierarchical system [START_REF] Kottak | The process of state formation in Madagascar[END_REF][START_REF] Raherisoanjato | Des territoires et des contrats au service de l'environnement. Le Lalangina, sur le versant ouest du COFAV (corridor forestier Ambositra Vondrozo) à Madagascar[END_REF]. Ambendrana (Site 1) was part of the Lalangina chiefdom [START_REF] Kottak | The process of state formation in Madagascar[END_REF],

and Sahabe (Site 2) belonged to the Arindrano chiefdom [START_REF] Giambrone | Teto anivon'ny riaka[END_REF][START_REF] Kottak | The process of state formation in Madagascar[END_REF]. After unifying the country during the 19 th century, the Merina reinforced drainage and irrigation to favor paddy rice cultivation and collected taxes. The French colonial government ruled Madagascar from 1896 to 1960.

Details on Ficus species found in the study area (Table 1) show aspects of their ecology, their life forms, fruit size and geographical distribution. We analyzed the uses and values of all the nine Ficus species found in our sites, and looked in detail only at the distribution of the three most common species, Ficus lutea, F. reflexa and F. tiliifolia. These species are found mainly in eastern and southeastern Madagascar at altitudes below 1700 m [START_REF] Dalecky | Moraceae, Ficus and associated fauna[END_REF][START_REF] Rafidison | Ethnobiologie et écologie des Ficus des terroirs Betsileo et du corridor Ranomafana -Andringitra (Madagascar). Dissertation University Antananarivo Rafidison[END_REF].

We conducted ethnobotanical studies during successive periods of 15 days totaling six months in the two sites. Open-ended interviews were conducted with 64 (Ambendrana) and 90 persons (Sahabe), children, adults and elders of both sexes from farmer families (Table 2). responses would be used for scientific purposes and all gave their consent. Heads of these villages and major informants were invited to Antananarivo in 2016 and contributed as authors to a paper presenting preliminary results [START_REF] Rafidison | Pratiques et connaissances naturalistes des communautés Betsileo: lisière du corridor forestier Andringitra-Ranomafana, Madagascar[END_REF].

We determined the distribution of individuals of the three most common Ficus species across the two rural landscapes according to their position relative to distinct land-use units distinguished by local people. Land-use units were defined as social-ecological facets, building on the concept of ecological facet used by [START_REF] Blanc-Pamard | Dialoguer avec le paysage ou comment l'espace écologique est vu et pratiqué par les communautés rurales des Hautes Terres malgaches[END_REF]. We define a social-ecological facet as a locally recognized spatial entity characterized by homogeneous ecological and use characteristics. All social-ecological facets are designated by distinctive names in Betsileo.

These names are interpreted locally to represent types of land associated with a particular set of practices. They include land-use units as well as elements such as tombs, megaliths, rivers, rivulets and paths that are markers of agrarian and forest territories (Table 3). We complemented the list of facets recorded by [START_REF] Blanc-Pamard | Ambendrana, un territoire d'entre deux : conversion et conservation de la forêt (Corridor Betsileo, Madagascar) Fianarantsoa, IRD/CNRE[END_REF] in Ambendrana and identified those of Sahabe.

The distribution of individuals of the three most common Ficus species in 31 different hills, each generally associated with one farm (18 in Ambendrana and 13 in Sahabe), was recorded with the help of owners of the land and other inhabitants. They provided ethnobotanical data during field surveys on all Ficus species. We identified all Ficus individuals to the species level and characterized the social-ecological facets in which the three most common species occurred and noted GPS coordinates. All elements of the landscape, such as houses, planted or spontaneous vegetation, rivers and cliffs were noted, and schematic drawings were made in order to obtain a visual record of the spatial relationships between the different elements of the rural landscape. An example of these schematic drawings is given in Fig 2.

Almost all Ficus individuals of the most common species found in the territories of the two villages were noted. Given the social importance of Ficus, local experts were able collectively to locate exhaustively all individuals of each Ficus species, except for those found in inaccessible areas such as cliffs. For example, for Ficus reflexa, all individuals planted around zebu corrals were counted. Similar exhaustive censuses were conducted on all three most common Ficus species.

We aimed at understanding population dynamics of F tiliifolia (the only species out of the three chosen for a detailed study that was not planted or very rarely), to identify whether this species could maintain itself without human interventions. We classified F. tiliifolia individuals into three categories on the basis of their architecture, following [START_REF] Hallé | Tropical trees and forests. An architectural analysis[END_REF], and functional categories of 'trees of the past' or old (trees with complex branching patterns, large round to flat crowns), 'trees of the present' or mature (trees with simple branching patterns and a pyramidal crown) and trees of the future or young (saplings with few lateral branches) [START_REF] Oldeman | Diagnosis of the temperament of tropical rain forest trees[END_REF]. These functional categories are supported by studies on tree architecture that consider repetitions and reiterations as elements of trees' ontologies [START_REF] Barthélémy | Plant architecture : A dynamic, multilevel and comprehensive approach to plant form, structure and ontogeny[END_REF]. Furthermore, we cross checked the trees' relative age with the help of elders' local expert knowledge.

.

RESULTS

We conducted ethnobotanical surveys of uses and practices related to the nine Ficus species found in these two sites with 154 informants. Distribution patterns were documented in detail for the three most abundant species. In the two sites, we recorded a total of 195 isolated trees of Ficus tiliifolia, 138 individuals of Ficus reflexa, isolated or in clusters in small woods or live hedges located around zebu corrals, and 29 individuals of Ficus lutea, isolated or mixed with F. reflexa around zebu corrals.

Naming and symbolic and material uses of Ficus species

All Ficus species are known locally, bear names shared by all inhabitants and have one or several associated symbolic and material uses; sets of uses differ among species (Table 4).

Naming

Names are symbols generally linked to significant meanings that are linked either to a characteristic (motivated) or are arbitrary (non-motivated). All nine Ficus species bear motivated names. The Betsileo have no generic taxon name that corresponds to the genus Ficus. But once people realized our interest for Ficus species, they showed without hesitation all Ficus species growing in their territory and only Ficus, even those we did not know initially.

Some names are based on a system of correspondence or analogy with the human body and other names refer to the social significance accorded to the tree, its morphology, the type of fruits it bears, the uses of its leaves and potentially the tree's links to other cultural groups, especially the Merina.

Each species bears a distinct motivated name except for two pairs of species that bear each the same motivated name (F. politoria and F. brachyclada; F. botryoides and F. trichoclada).

The first two, like many species of the section Sycidium, have scabrous leaves that can be used as sandpaper and are named kivozo or ampaly (literally, "scabrous"). The second pair, named fompoha, have young tree stages that are similar morphologically and are closely related species in section Sycomorus. The name amonta (literally, "abundance") is given to Ficus lutea, because it is associated with noble Betsileo classes (hova). Seedlings are protected for this reason. "Avelao hitsiry ny amonta dia ny taranaka sy ny harena mba hamontafonta" (Leaving the seedling of amonta to grow will make local descendants also rich). Although seedlings are allowed to establish anywhere, poor people are forbidden from planting this tree near their houses.

The name nonoka, applied to F. reflexa, means a feeding breast or the act of sucking milk (Table 4). The abundant white latex produced when the tree is wounded is associated with human milk and by analogy, leaves are used by breast-feeding women to increase milk production.

Voara (F. tiliifolia), (literally "fruit of ara", the term ara meaning literally, "what is linked to" presumably to all people) is said to be the "reninkazo" (literally mother tree) and is a symbol of fecundity. People relate this name to the tree's height and its abundant fruits, appreciated by children and zebus, its uses for making clothes and as a medicine for women giving birth. Voara is perceived as a tree that attracts and maintains water in the soil. By analogy, cutting a voara is a negative act that may lead to the drying up of children's tears or of a mother's milk. Two varieties of Voara are named according to the distinctive traits of their fruits, such as sweetness (Voaramalefaka) or an atypical shape of the fruit (Voarabekobo, meaning "with a thick lip").

Ficus trichopoda is named aviavy meaning in Betsileo "arriving", the same name is used by the Merina for Ficus polita in Antananarivo. Tsaramady, used by the Betsileo for F. polita, means literally "the good one".

Uses

Uses of fig trees may vary according to the socio-ecological facets in which they grow. These uses, both symbolic and material, are dependent on whether the trees grow naturally or have been planted. Meanings and roles of the different Ficus species also vary depending on their historical connections with the Betsileo. For instance, the social significance of F. tiliifolia is strongly influenced by the past use of its bark as a textile, a use that was renewed during the turmoil of the 1940s for making blankets or coats which had a high economic value; a coat could be exchanged for a zebu.

Material and symbolic medicinal uses

All Ficus species have medicinal uses for the Betsileo and all are used today to treat diverse diseases (diarrhea, cough, fever, worm infections) and to treat wounds and a range of complaints related to women's health. Some uses are shared between two or more species. The latex of F. lutea, F. tiliifolia and F. reflexa is used to heal wounds. Healing practices used by ombiasy (healers) make use of all Ficus species found in the area. Ancestors' spirits, called upon by the ombiasy, are said to systematically designate Ficus products as important remedies.

The ombiasy receive this information through dreams or during possession rituals.

Other material and symbolic uses

All Ficus species have technical uses that confer to them an economic value. Some species are widely recognized by the Betsileo to be consumed by animals such as birds, bats and lemurs, but some are known to be particularly attractive (F. reflexa and F. lutea). This knowledge is used when Betsileo hunt bats. The latex of F. reflexa and F. trichopoda is used as birdlime.

Fruits of F. tiliifolia, F. lutea, F. reflexa and F. botryoides are eaten by people, mainly by children. The fruits of F. tiliifolia were traded before new fruit species were introduced into the area.

The bark of both F. politoria and F. pachyclada is used for making strings and ropes, while the bark of F. tiliifolia and F. botryoides was used for making textiles and is symbolically associated with past lives of ancestors. Other technical uses are numerous, including planting of stem cuttings to prevent erosion through making living hedges (F. reflexa and F. botryoides)

and use of dry leaves as fertilizer (F. tiliifolia) (Table 4). All these species are used as land markers but the species most frequently used for marking ancestors' tombs are F. reflexa and F. lutea.

Tree management practices: an ethnohistorical construction of Betsileo landscapes

All Ficus species have symbolic and material uses that together determine when and where they will be protected or not. Symbolic uses of a given species can vary depending on the social-ecological facets in which individual trees grow. Indeed, any Ficus tree that grows The ombiasy empower the tree through magical practices by adding a charm to trees planted on the border of zebu corrals to protect the latter against evil spirits or robbery, a practice named "tafotombala" (literally: "that brings wealth").

Reading the landscape: ethno-historical indicators

Records of oral memory of the Betsileo, together with their local knowledge of landscape elements, show that their indicators of the composition of their rural landscapes are socialecological facets, encompassing material (e.g., ecological and topographic aspects, types of agricultural practices or human habitats) and symbolic aspects (e.g. burial places, other markers of the memory of ancestors). Naming of social-ecological facets is precise (Table 3). They can be grouped into five large categories: vegetation types, agricultural land-use types, topographical elements, extraordinary or specific abiotic facets, and historical and sacred facets.

Based on these indicators, we identified three overlapping ethno-historical zones along an east-west axis in site 1-Ambedrana (Zone A-eastern forest; zone B-protected hill-tops; Zone C- We do not have an explanation for the absence of a Zone B-protected hilltop in this site. We visited areas near forest fringes where none of the three Ficus species most common in our two sites were found.

Management practices and distribution patterns of Ficus trees

Two groups of Ficus species can be differentiated according to how individuals establish within the rural landscape. Species of the first group regenerate exclusively through self-sown seedlings (F. tiliifolia, F. politoria and F. brachyclada). Species of the second group reproduce by seedlings but are also actively propagated by large pole-like cuttings (F. reflexa, F. lutea, F.

botryoides, F. trichoclada, F. polita and F. trichopoda). We recorded in situ only individuals of F. trichopoda that were propagated by cuttings (six individuals in site 1 and site 2 in Zone C-15 roofs area far from the forest), but local experts explain that seedlings exist, are rare and are protected (Table 2). Ficus trichopoda is found growing in the forest only on the eastern side of the mountain ridge. Such trees were incorporated into agricultural lands progressively as fields became permanent.

Regeneration only by seeds: example of F. tiliifolia

According to local knowledge, all Ficus tiliifolia found across the landscape were trees maintained mostly by forefathers due to their multiple values. F. tiliifolia very rarely regenerates now in agricultural lands. It is still generally protected as a part of the heritage but in some cases, when the tree affects crop productions, individual tree may be cut. It is otherwise maintained for its qualities for fertilizing soils and for maintaining soil water, when it does not hinder agricultural production (Table 4). Analysis of the population structure of F. tiliifolia in the two study sites shows that old trees are predominant (91%) with only one individual representing the young stages (Fig 6).

Ficus species that both regenerate by seeds and are propagated by stem cuttings: examples of Ficus reflexa and Ficus lutea

Ficus reflexa is rare in Zone A-eastern forest areas in both Ambendrana According to ethno-historical accounts, they were planted by noble Betsileo classes, as a symbol of their power over land and a sign of their connection to the Merina kingdom.

Individuals growing spontaneously on tombs are strictly protected. Some individuals are found regenerating naturally at the foot of the cliffs both in Ambendrana (few) and Sahabe (numerous, not counted).

DISCUSSION

Of the 24 Ficus species identified in the forest corridor that joins Ranomafana and Andringitra National Park (Rafidison et al 2015), nine are present and protected as isolated trees or as clusters in live hedges or in small woods in the adjoining rural landscapes of Ambendrana and Sahabe. [START_REF] Martin | Conservation value for birds of traditionally managed isolated trees in an agricultural landscape of Madagascar[END_REF], show that 33 % of isolated individuals (sample size, 48) of tree species are represented by three Ficus species (F lutea, tiliifolia and trichopoda).

The protection of the nine Ficus species is driven by their multiple uses and varies depending on their distribution in social-ecological facets. Ethnohistorical accounts show temporal continuity of some of the practices associated with these species, and also document changes over time. Distinct sets of uses (past, present, symbolic and material), are associated with each species. Multiple relational, economic and ecological values link uses and the distribution of Ficus trees in the landscape following social-ecological dynamics. Distribution patterns of the three species show that they are predominantly protected as "frontier" elements between social-ecological facets. Local people explain the greater numbers of these trees at the edges of agricultural lands by the fact that they are cut within agricultural areas when they hinder production, but are left in other places to fertilize soil and maintain soil water supply.

Our analysis of the protection of Ficus trees also shows links between symbolic, material and economic dimensions. For example, when found near zebu corrals, Ficus trees play symbolic roles through charms to protect zebus and humans, but also have an important role in preventing soil erosion, thereby maintaining the sunken basins cut into the soil that serve as zebu corrals.

No strict taboo (fady) enforces their protection, but some sayings encourage people to protect them. However, a fady or strict rule forbids the active plantation of Ficus lutea and F. trichopoda, especially by ordinary people, although plants of these species that originate from self-sown seeds can be protected. Plants that establish naturally from seeds are considered by local people to have been planted by ancestors and other intangible creatures, and are highly valued, especially when they grow in social-ecological facets such as burial areas, steles or other landscape features such as rocks and cliffs. Out of the nine species, seven are actively planted by stem cuttings (Table 4) following decisions about where they should be planted and choice of social-ecological facet(s). Regarding the species that regenerate exclusively by seedlings (rare), their degree of protection remains unclear because they are rare.

Vernacular and scientific taxonomy

Our results show that identification and a comprehensive naming system of Ficus species is shared by all inhabitants from elders to children in both sites. This enhances the capacity of the Betsileo as a cultural group to share knowledge and rules pertaining to management practices, a mechanism known from many traditional societies (e.g. [START_REF] Friedberg | Classifications populaires des plantes et modes de connaissances[END_REF][START_REF] Ellen | The cognitive geometry of nature: a contextual approach[END_REF]. How Betsileo identify Ficus species and how scientists identify them are quite similar.

Although they do not use a generic category encompassing all Ficus, the Betsileo identify Ficus species by observing similarities and differences among them in morphological and other traits of leaves, fruits, bark and latex. Differences in growth habits, such as those between hemiepiphytes or lithophytes (able to germinate and grow on rocks) and those that grow only as standing trees, are also considered. Aspects of their phenology, such as the asynchronous production of leaves of some Ficus species, are also taken into account. The traits used to identify different species are often those associated with the uses of each species. For instance, the two species found locally whose leaves are scabrous are both named ampaly, meaning scabrous (Table 4), and are among the few pairs of species that bear the same name. They are in fact sister species in the section Sycidium of the genus (Table 1, 4).

The Betsileo eat the fruits of Ficus tiliifolia, F. botryoides and F. trichoclada. These related species all belong to the section Sycomorus of the genus (Table 1). The large and conspicuous crops of figs produced by these species are well-known by scientists to attract mammals [START_REF] Shanahan | Fig-eating by vertebrate frugivores: a global review[END_REF][START_REF] Harrison | Seventy-seven ways to be a fig: an overview of adiverse assemblage offigs in Borneo[END_REF]. Fruits of many species of section Sycomorus are widely eaten by humans, for example in Africa [START_REF] Burrows | Figs of Southern and South Central Africa[END_REF].

Ficus botryoides and F. trichoclada bear the same local name (fompoha) and share the same uses, and indeed are sister species in the phylogeny of Ficus. Both are cauliflorous (bearing fruits on the trunk). Both grow in riverine forests and have very similar leaves (F. Kjellberg, unpublished data). Ficus sycomorus, the type species of section Sycomorus, which produces prolific amounts of fruits, is protected and its fruits are eaten near Tulear in western Madagascar (V. Rafidison, unpublished data). This species is widely utilized throughout Africa for its fruits, including in Egypt, beyond its natural distribution area, where it was domesticated by the ancient Egyptians [START_REF] Burrows | Figs of Southern and South Central Africa[END_REF].

Other Ficus whose fruits are eaten are F. lutea, F. trichopoda, F. polita and F. reflexa.

These species belong to section Galoglychia. The fruits of the first three species are consumed by humans because of their large size. In the area of Antananarivo, for example, the unripe fruits of F. polita are traded and consumed for their well-known benefits for the throat (Aumeeruddy-Thomas et al 2018).

A most fascinating characteristic of species of section Galoglychia is their hemi-epiphytic and hemi-epilithic growth form. This characteristic, associated by the Betsileo with the idea that the seeds are "planted" by intangible creatures, is the reason why people protect them especially on sacred social-ecological facets, such as rocky steles or megaliths, and in small isolated woodland patches (songonata) within the open landscape. These woodland patches derive from nucleation processes as explained above [START_REF] Martin | Conservation value for birds of traditionally managed isolated trees in an agricultural landscape of Madagascar[END_REF]Rafidison, 2013). Such vegetation dynamics are also known for 'orphan' trees protected in swidden agriculture in Africa [START_REF] Carrière | Orphan trees of the forest. Why do Ntumu farmers of southern Cameroon protect trees in their swidden fields[END_REF] and in Asia [START_REF] Eden | Isolated Ficus trees deliver dual conservation and development benefits in a rural landscape[END_REF]. These small woods within open Betsileo landscapes acquire a sacred value, being associated with tombs or steles and Ficus species.

Most of the uses of Ficus documented in the Betsileo sites, such as their use to improve milk production by breast-feeding women, or by cattle, or their use as live hedges for protecting people or cattle, are similar to those of the same species elsewhere in Madagascar or of other Ficus species in Africa (e.g. [START_REF] Burrows | Figs of Southern and South Central Africa[END_REF], Asia (Kunwari and Bussman 2006) and the Pacific [START_REF] Walter | Fruits d'Océanie[END_REF].

Continuities between past and present uses, symbolic and material

Continuity between past and present-day uses is a key driver of the protection of isolated

Reciprocal relations, tree agency and interconnected multiple valuing systems

Trees are at the heart of exchanges between humans, other living beings (biodiversity) and intangible entities [START_REF] Aumeeruddy-Thomas | Agroforestry[END_REF]. The roles of isolated or clustered Ficus individuals as mediators between people and ancestors and other intangible entities, as well as between people, their landscape history and their identity, and between people and the animals they hunt or that are perceived as disseminators of seeds (e.g., birds and bats), makes these trees active hubs. The Betsileo do not conceive of Ficus trees as simply passive elements, but as recipients of social actions built upon reciprocal relations between the Betsileo and the power of nature, designated in Madagascar as hasina or sacred power, which is likened to God and which can be transmitted by people or plants, as shown for F. lutea in Antananarivo (Aumeeruddy-Thomas et al. 2018). To the Betsileo, trees have a level of agency, because they directly affect people's lives by bringing happiness or protection, or by purifying people of their deeds. They therefore also affect social organization and decisions, and in that respect form part of the hybrid communities that link humans to other living beings, as is true for many other

Tree management and propagation practices and distribution

Naming and uses are similar or confounded between the Betsileo and the Merina cultural groups who live in the region of Antananarivo, where we conducted a previous study. In that study we focused on F. lutea in the sacred hills of Antananarivo, and we showed that the Merina used many symbols to portray their supremacy, among them F. lutea. Trees of this species were planted by stem cuttings in new places they had 'acquired' during their unification of Madagascar as a state (Aumeeruddy-Thomas et al. 2018). Similar practices have been described

in continental Africa, where Ficus trees were planted in front of conquerors' houses or on their tombs to mark their acquisition of a new territory [START_REF] Dury | Ethnobotanical approach to Ficus in northern Cameroon, Master's Dissertation UNESCO/Man and the Biosphere Program[END_REF]. The use of the name amonta for polita in Antananarivo (aviavy). This pattern also applies to F. reflexa, which bears the same name (nonoka) in Betsileo and Merina. The above species are considered to have the power to protect humans and zebus, to purify people, or to bring happiness (but also death). These beliefs all portray the supernatural power accorded to them and a strong influence of the powerful Merina.

Potential and constraints of Ficus tree protection

The diverse distribution patterns of F. tiliifolia, F. reflexa and F. lutea show very distinct dynamics that are taken into account in efforts of local people to protect them.

Ficus tiliifolia

While F. tiliifolia is accorded a positive value in agriculture for its role in fertilizing soils, it can also negatively affect agricultural production. The tree is deciduous (i. analogies between people and trees ;

(3) its role as food for children and cattle; (4) its role as an identity marker for Betsileo people. In Sahabe, where F. tiliifolia is rare, people pay less attention to this tree than in Ambendrana. A possible explanation is that Sahabe is located at a higher altitude and the climate is colder and less suitable for this species. Furthermore, the rural landscape in Sahabe shows a larger extension of herbaceous grasslands associated with a higher level of pastoral activities (Fig 4). As explained by [START_REF] Kottak | The process of state formation in Madagascar[END_REF], the Arindrano chiefdom was probably more based on pastoralism than the Lalaina kingdom. Virtually restricted to forest environments, F. tiliifolia could not survive in the savannah-type environments of Sahabe (just as its population is now shrinking in the open areas created by sedentarized agriculture in Ambendrana).

Ficus reflexa

Ficus reflexa is protected owing to its multiple uses. It is abundant in a large area of plains and near the cliffs, areas that were occupied by the Betsileo before periods of insecurity in the 17th and 18th centuries, and then occupied by the Merina in the 19th century, the area defined in this study as Zone C-15 roofs area. The distribution of F. reflexa in this zone is thus the result of historical accumulations of trees of this species planted by people, reflected in its abundance in abandoned villages (tanana haola) and in zebu corrals, both abandoned ones and those presently used.

Ficus lutea

Ficus lutea is also protected in Zone C-15 roofs area, but is less actively planted due to its association with noble classes. It is therefore much less common than F. reflexa, although it has multiple uses.

Implications for conservation and potential collaborative approaches

Conservation efforts increasingly consider anthropogenic landscapes made of the co-existence of agroecosystems and other elements such as forests, pastures, fallow land, that contribute to connectivity and habitat diversity [START_REF] Perfecto | The agroecological matrix as alternative to the landsparing / agriculture intensification model[END_REF][START_REF] Manning | Transferring biodiversity-ecosystem function research to the management of 'real-world' ecosystems[END_REF].

Isolated and clustered Ficus individuals are key elements of Betsileo rural landscapes. Given the multiple economic, relational and ecological values associated with them, Ficus trees can contribute both to biodiversity conservation and to support social-ecological systems built on local knowledge. "Community-based" approaches that were developed previously in Madagascar-and more widely across the planet-have not led to the expected results, (Ward et al. 2014, Blanc-Pamard andFouroux 2004). This suggests that multiple-evidence and collaborative approaches that build on synergies between local and scientific knowledge to understand multiple valuing systems should be further implemented [START_REF] Pascual | Valuing nature's contributions to people: the IPBES approach[END_REF], Tengo et al. 2014) protection also due to respect for ancestors and multiple uses. wood used as a purifier if a fady (taboo) is not followed (S) natural reproduction by seed and planted by cuttings this tree is never planted, but rare seedlings are protected du to the benefits the tree brings. Table 1 Ficus species found in the study areas, their major biological and ecological characteristics, habitats and geographical distribution

  Fompoha (literally "to blow") relates to the small cloud of tiny insects that come out of the ostiole (opening at the apex of all figs, through which fig wasps penetrate to pollinate the flowers inside, and through which their offspring emerge when the fruit matures) when one blows on a mature fig. Children readily demonstrated to us how they blow on F. botryoides fruits to make the fruit more palatable.

  from self-sown seedlings near social-ecological facets such as tombs, steles, abandoned ancient villages or elements of landscapes such as large rocks, are systematically protected. Indeed, their seeds are perceived to have been brought into these places by ancestors or other intangible creatures. Fig trees are therefore a cultural element of the landscape and not solely an element of natural regeneration. When growing in the middle of a field, where they may hinder agricultural production, spontaneously establishing fig trees of all species can be cut, despite a number of sayings and beliefs that encourage their protection. There is no strict fady (taboo) that protects Ficus species from being destroyed. Ombiasy also indicate to local inhabitants how to plant particular Ficus species.

  15 roofs area) (Fig 3). The "15 roofs period" refers to the period, during the 19 th century, when Merina administrators developed larger villages for the Betsileo, associated with markets closer to the lowlands. In Zone C-15 roofs area, the landscape is shaped both by activities conducted before the unification of Madagascar by the Merina and those developed with the establishment of a new village center during the Merina period, and contemporary periods. Details of the landscape composition including traces of past activities, vegetation types, agricultural practices and Ficus distribution are given in the online resources (Online resource 1). In Sahabe, pasture lands occupy larger areas than in Ambendrana; forest remnants are few and pasture and cultivated lands form the major part of the rural landscape. Small areas of forest fallows, associated with past tavy practices, are still found within Sahabe's open rural landscape. We characterized the rural landscape in Sahabe as a Zone C-15 roofs area for the purpose of comparison with a similar ethnohistorical area in Ambendrana. No east-west axis is visible, in contrast to Ambendrana (Fig 4).

Ficus

  tiliifolia is abundant (70 individuals) in Zone A-eastern forest in Ambedrana but absent in Sahabe. In Zone B-protected hill-tops (this zone is inexistent in Sahabe), F. tiliifolia is less abundant (21 individuals) in Ambendrana than in Zone C-15 roofs area, with 65 individuals in Ambendrana and 39 in Sahabe. In Ambendrana, F. tiliifolia individuals are located at the edges between different social-ecological facets, marking transitions between two types of landscape elements. Such positions are occupied by 57 % of 70 individuals in Zone A-eastern forest, 81 % of 21 individuals in Zone B-protected hill-tops, and 59 % of 65 individuals in Zone C-15-roofs area In Sahabe, individuals located between two social-ecological facets, in Zone C-15 roofs area represent 41% of 39 individuals and are totally absent near the eastern forest fringes (Online resource 2). The abundance of F. tiliifolia between two social-ecological facets indicates that they have been cut within each facet, e.g., during tavy clearing activities. The photo shown in Fig 5 portrays an individual originating from the forest that has persisted on the edge of a paddy field.

  (three individuals) and in forest fringes of Sahabe (five individuals). In Zone B-protected hilltops, F reflexa is absent in Ambendrana. It is most abundant in site C-15 roofs area in both sites, with In both sites, F. reflexa is mainly located at interfaces between two social-ecological facets (71 % of 44 individuals at Ambendrana; 93 % of all individuals at Sahabe). They are associated with vatobe (steles) and vatolahy (very large ancient megaliths), fasana (sacred woods), alagasy (forest), and tanimboakazo (orchards), places where they establish as seedlings and are then protected. Valanomby (zebu corrals), tananahaolo (abandoned villages), cour (courtyards), and valamparihy (mud walls along rice fields) are typically facets where individuals of this species are planted as pole cuttings. Some individuals are associated with tombs, thickets, pasturelands and threshing areas or grow spontaneously as hemiepiphytes on old forest trees bordering fields or as lithophytes on rocks (Online resource 3) Ficus lutea is rare in Ambendrana (five individuals) and found only in zone C-15 roofs area. Several plants from self-sown seedlings were found nearby on the cliffs (not counted). In Sahabe, this species is more abundant (24 individuals). In this site, most individuals are located at edges between two social-ecological facets (71 % of 24 individuals). They grow from spontaneous seedlings, which are always protected. They are also planted by large stem cuttings, but only members of noble classes are allowed to plant them, never ordinary people.They are absent in Zone A-eastern forest and Zone B-hill tops. Some individuals are also found on the humid eastern slopes (Tanala side) of the mountain ridge (V.Rafidison & F. Kjellberg, unpublished data) (Online Resource 4).According to local knowledge in both sites, plants of F. lutea growing in Zone C-15 roofs area originate from seeds brought by birds and bats that visit the cliffs or orchards. Individuals found in abandoned villages (tanana haola) in courtyards of noble people or rayamandreny and in old herbaceous fallows (songonala), originate from the plantating of large stem cuttings.

Ficus

  trees, or clusters of them, found in Betsileo rural landscapes. Human memory associated with transmission of local knowledge by forefathers, and ways people view this transmission as part of their identity, are key to linking people to biodiversity (including agrobiodiversity), places and landscapes (Nazarea 1999). Roles of trees in building territories and relationships to nature have been highlighted in many societies. Their perennial dimensions, multiple economic uses, and perceived symbolic analogies linking humans to trees are characteristics that favor their presence in many local territories. Their protection is further enhanced by diverse beliefs in their roles as mediators, or as hosts of intangible entities that may have positive or negative impacts on human well-being(Aumeeruddy and Bakels 1993;[START_REF] Rival | The social life of trees: anthropological perspectives on tree symbolism[END_REF] Aumeeruddy- Thomas et al. 2018).Trees, in this case Ficus species as perennial species, are transgenerational markers of local social-ecological systems and landscapes and link the present to cultural memory. Their role in the memory of the Betsileo can be compared to that of other permanent features such as topographic elements (hills, mountains, cliffs or rocks). Links between their symbolic and practical uses and the trees' ecology, their spontaneous regeneration by seeds in specific habitats, or their propagation by stem cuttings, are jointly used for their management, as shown also for F. carica in the Mediterranean region[START_REF] Aumeeruddy-Thomas | Fig and olive domestication in the Rif, Northern Morocco. Entangled human and tree lives and history[END_REF].Isolated or clustered individuals ofFicus within open agricultural landscapes participate in the construction of these rural landscapes, as markers of local history and as testimony to local attachment to the land and its resources. This attachment contributes to reinforcing local governance of social-ecological systems. During times of turmoil faced at the national level when threats on biodiversity, agroecosystems and local livelihoods are increasing, looking at ways local knowledge is linked to remarkable trees from cultural, economic and ecological perspectives, may help identify sustainable solutions to reinforce local social-ecological systems and livelihoods. Understanding such systems requires building bridges between local knowledge and scientific knowledge. A study conducted by Marie et al. (2009) on local practices related to an endangered group of tree species in Madagascar, the seven endemic baobab species (Adansonia spp.), shows that conservation narratives about threats relating to Malagasy baobab often attribute these threats to human activities, but neglect consideration of how baobabs are part of local agroecosystems and are often protected, if not cultivated, by local farmers.

  indigenous peoples and local communities across the planet[START_REF] Stepanoff | Hybrid Communities: Biosocial Approaches to Domestication and Other Transspecies Relationships[END_REF].[START_REF] Diver | Recognizing "reciprocal relations" to restore community access to land and water[END_REF] argue that in Madagascar, taking into account reciprocal relations between humans and biodiversity is crucial to improve conservation approaches. In particular, their study shows that local rules of access to resources and places reinforce reciprocal relations. In our study sites, the distribution of Ficus species across the landscape does not follow haphazard patterns, and associations of different Ficus species with different socio-ecological facets, i.e., specific places having strong meanings, leads to formulation of rules that guide access to and management of these places and the Ficus they include. These rules have to be considered in collaborative approaches for biodiversity conservation if local livelihoods are to be sustained.Past and present uses of biodiversity, transmission of knowledge and local formal and informal rules guiding access and management approaches to resources contribute to enhance relational values that link people to landscapes[START_REF] Chan | Why protect nature? Rethinking values and the environment[END_REF]. This needs full consideration if conservation efforts are truly to engage with indigenous peoples and local communities (Diaz et al. 2018). In addition to their strong social or relational value, Ficus trees also have an ecological value as ecosystem engineers in sacred woodland patches in open agricultural areas in Betsileo rural landscapes. Their economic value is related to their roles in checking erosion and other multiple material uses. An interconnected multiple valuing system characterizes the use of isolated and clustered of Ficus trees in the Betsileo rural landscapes. As discussed by Pascual et al. (2017), multiple valuing systems are crucial to understand and sustain to attain sustainable development objectives.

F

  . lutea by the Betsileo is very similar to its Merina name amontana. Both in Antananarivo and in the Betsileo sites, F. lutea is associated with similar sets of symbolic values. All this suggests a strong Merina influence. In the rural landscape, F. lutea is mainly planted from stem cuttings by noble classes. It regenerates from seed only in the cliffs area (Zone C), where ancient hova (Betsileo nobles) lived[START_REF] Moreau | Les gens de la lisière[END_REF]. The absence of F. lutea in Zone A-Eastern forest (except in the humid Tanala side located on the eastern slope of the mountain ridge) suggests that it may have been brought into the Betsileo region by the Merina. The same pattern applies to F. trichopoda, which in the Betsileo study sites occurs only (except for rare seedlings) as planted individuals propagated from stem cuttings. This species bears the same name as F.

  e., leaves of an individual are shed synchronously) but individual trees differ in when leaves are shed. Some individuals shed leaves at a time when the falling leaves can harm young rice seedlings. Farmers may decide to cut such trees. Martin et al. (2009) suggested that protection of Ficus species by the Betsileo was linked to respect for ancestors' practices, and roles of taboos (fady). The present study corroborates these findings and revises ideas about taboos. Taboos are often considered instrumental in reinforcing conservation but they can be lifted (Fernandez-Llamazares et al. 2018). Ethnobotanical and ethnohistorical approaches taken in our study show that several factors-historical, symbolic, but also economic-contribute to explaining why, what and where Ficus trees are protected in Betsileo rural landscapes. Patterns of protection are not linked to any strict taboo. Although people respect the practices of ancestors and refer to this in the first place, the Betsileo give numerous other reasons to explain why they have protected F. tiliifolia trees. Four major and interlinked reasons are given: (1) the numerous practical uses of this species, e.g. fertilizing soils and improving soil water content; (2)

  biodiversity and sustain local social-ecological systems. The following steps are some approaches that we suggest:-Characterizing rural landscapes and their complexities based on i) local knowledge and other tools (e.g. remote sensing, ecological analysis, naturalists' knowledge of biodiversity) and ii) a fine comprehension of social-ecological facets, distribution of practices and changes in practices in the landscape, over time; -Identifying key elements such as Ficus trees through multiple evidence-based approaches based on local knowledge and joint investigations of ecological, ethnobotanical and anthropological aspects;-Defining together through focus-group discussions with elders, men, women, and children, the key roles of specific species as indicators of well-being and their diverse values for people and how they affect other groups of species. Discussing potential plans for supporting the protection of such species, with due recognition of values, of who is entitled to plant each species, and how and where (in which specific social-ecological facets) each can be propagated; -Avoiding inventing "local" institutions that are not truly appropriated by local people, and building a joint understanding of collective visions and rules of access to resources.-Constraints in such approaches lie in the different values accorded to biodiversity by conservation managers, scientists and local communities and the necessity to identify parts of their interest that can be shared as a first step for developing learning processes CONCLUSION The roles of trees in live hedges, in small woodland patches or as isolated elements in rural landscapes are known, especially in agroforestry systems. However, the multiple social, historical and political roles of trees in open landscapes have rarely been discussed in relation to collaborative conservation approaches that consider local social-ecological dynamics. This case study provides an understanding of how the Betsileo people associate multiple relational, economic and ecological values with Ficus trees at the landscape level. The ecological role of Ficus trees as stepping-stones are known to enhance connectivity and facilitate re-afforestation in fragmented rural landscapes. They may be considered for their potential to improve the management of biodiversity and for their multiple local values . Propagation techniques applied to each species, and the places where trees of each species are allowed to grow, are linked to interconnected cultural, ecological, historical and economic contexts. This study suggests avenues regarding approaches to understanding the multiple values of trees in open rural landscapes and their potential roles for supporting conservation of local social-ecological systems outside protected areas.

  ancestors (S) ; presence linked to divinities and intangible creatures (S) fallen leaves fertilize soils (M) roots maintain water in the soil (M/ S) bark used for making a cloth named ato (M,)-Previously a coat made with fato could be exchanged against a zebu (M/S) the fruit is eaten. Two varieties known : voara bekoba and voaramalefaka (M) leaves have medicinal uses : heal stomach ache, skin disease, facilitates birth (M,S) natural reproduction by seed, seedlings sometimes transplanted local sayings encourage people not to cut this tree because this would have negative impacts on soil water, and would dry up children's tears and the milk of breast-feeding women.

Fig. 1 Fig. 2 Fig. 3 Fig. 4 Fig. 5 Fig. 6

 123456 Fig.1 Location of the two study sites Ambendrana and SahabeRanomafana and Andringitra National Parks and the forest corridor, Madagascar

  restricted access shaped solely by biological considerations[START_REF] Marie | Taking into account local practices and indigenous knowledge in an emergency conservation context in Madagascar[END_REF][START_REF] Diver | Recognizing "reciprocal relations" to restore community access to land and water[END_REF].Indeed, conservation policies in Madagascar are based on protected areas from which people are excluded. These policies, initiated by the French colonial government, have continued to the present, with investment from numerous international conservation NGOs. The network of protected areas was expanded in 2003. Additions included the establishment of ecological corridors such as the Ambositra-Vondrozo forest strip connecting Ranomafana and Andringitra National Parks that was created in 2008. This corridor, which borders our study

practices

[START_REF] Blanc-Pamard | Dialoguer avec le paysage ou comment l'espace écologique est vu et pratiqué par les communautés rurales des Hautes Terres malgaches[END_REF][START_REF] Blanc-Pamard | Normes environnementales, transferts de gestion et recompositions territoriales en pays betsileo (Madagascar). La gestion contractualisée des forêts[END_REF]

. Formal and informal social hierarchies, local institutions and practices (including rituals, such as zebu sacrifices in the case of Madagascar) and exchange networks define people's relationships within social-ecological systems

(Berkes and Folke, 1998)

. Such complexity is rarely taken into account in conservation policies in Madagascar, which are historically based on zonation 1 The Merina is the dominant cultural group that livesin the northern part of the Highlands in Antananarivo and in the surroundings of what came to be the capital city of the country. and area, acquired an official status as a "New Protected Area" with restricted access

[START_REF] Blanc-Pamard | Normes environnementales, transferts de gestion et recompositions territoriales en pays betsileo (Madagascar). La gestion contractualisée des forêts[END_REF] Rakoto et al. 2014

).

  Among all persons interviewed, the elders ray aman-dreny, (literally "father and mother" representing customary heads), and ombiasy (traditional healers), and lineage heads (adults or elders), had extensive knowledge of local history and social practices. Local ombiasy use Ficus species, among other plants, to mediate, through dreams and visions, interactions between humans, ancestors and intangible entities of nature[START_REF] Ph | La place et les pratiques des devins-guérisseurs dans le Sud-Est de Madagascar[END_REF]. Lineage heads and elders accompanied us to locate Ficus individuals and provided ethno-historical information on present-day practices and oral memory of past uses and practices. We recorded local names, sayings and stories relating to plants and places. All interviews were in Betsileo and were

	transcribed	literally.	Following	the	terms	of	our	research	permit

(No°:193/08/MEFT/SG/DGEF/DSAP/SSE-01/07/08), all informants were informed that their

  . Practical collaborative conservation work requires long-term involvement of scientists, conservation managers and local communities (e.g. Lama et al. 2001: Diver et al. 2019). Our engagement in collaborative approaches started with the co-writing of a paper with local experts (Rafidison et al. 2016). Based on our experience, and on findings by other authors about the roles played by trees in open agricultural landscapes in Madagascar (e.g. Marie et al. 2009), we suggest a few rules of thumb for developing collaborative approaches that conserve

Table

  Click here to access/download;Table;Table 1.rtf

							Evergreen
	F. polita Vahl Site	m	Hemi-epiphytic Ambendrana	15(40)	C Sahabe	4F	(humid and gallery) forest	600-700	M (all island?),CA
	Source: Dalecky et al. 2003, revised 2010 by F. Kjellberg, V. Rafidison. and Y. Aumeeruddy-Thomas Men 45 59
	d:dioecious; m: monoecious; R: ramiflorous; C: cauliflorous; A: axillary
	D: dry, F: fresh Women		19		31
	M (E, C, N, W, S): Madagascar (east, center, north, west, south), Co: Comoro islands; Se: Seychelelles; CA: Continental Africa; Me: Mascarenes; Al: Aldabra
	Total		64		90
	Precision on informants ages
	Children < 12years		11		12
	Youngsters 12-18 years		1		4
	Young parent 20-40 years		29		52
	Old persons		23		22

Table

  Click here to access/download;Table;Table 2.docx

	Type	Terminology	Signification
	Natural or anthropogenic	Tapoka	Marshy areas not yet transformed into
	vegetation types		paddy fields
		Alagasy	Literally, "malagasy forest", stands for
			non-disturbed forests
		Alakininina	Eucalypt forest
		Atikifafa	Literally, "within the herbaceous
			savanna", an area that was never
			cultivated but has been burned for
			pasture lands
		Kapoka	Post-agricultural regrowth or forest or
		(Ambendrana)	bushy vegetation fallows
		Popoka (Sahabe)
		Kilanjy	Worn out land with a fallow of
			herbaceous for an undetermined period
		Kirihitra	Bushy vegetation
		Songonala	Literally "forest tuft" meaning an
			isolated small wood.
	Agricultural land use	Dobo	Fish pond
		Kipahy	Cultivated terraces, most often irrigated
			paddy fields
		Lalankely	Path
		Tanimboankazo	Literally "land of fruit trees" meaning
			orchards
		Tanimboly	Literally "cultivated land": the field
		Tsihintany	Literally, « land mat » : threshing
			ground
		Valamparihy	Mud walls protecting rice fields
		Valanomby	Zebu stockyard
	Topographic element	Amorotambina	The lower border of hill slopes
			generally located just above paddy
			fields
		Lohasaha	The head of the field or valley
		Tambina	The lower part of hill slopes
	Extraordinary or specific	Hara	Rocky cliffs
	natural element	(Harambato)
		Renirano	Literally "the mother of the wate"r : a
			river
		Riandrano	A rivulet
		Vatobe	Big rock
	Historical site	Aritsa	Sacred wood where utensils and
			garments that have been used during
			burial ceremonies are thrown away
		Fasana	Tombs

Table

  Click here to access/download;Table;Table 3.docx

	Tanana Haolo	Literally, abandoned hamlet or dead
	or Valamaty	hamlet.
	Vatolahy	Literally, Male stone Stela or megalith
		which can reach 1 to three meter high
		and which has a commemorative
		function..

Table

  Click here to access/download;Table;Table 4.docx tree is a marker of territories(M) tree important in open areas for its shade (M)

	wood for previous domestic uses : for making
	dishes, spoons, and preparing baskets and
	container for feeding livestock and chickens
	(M)
	latex used as birdlime to catch small birds,
	especially the fody (Foudia madagascariensis)
	or to repair holes in jugs and buckets (M)
	Tree, symbol of fertility (S)
	F.botryoides
	Baker

Table

  Click here to access/download;Table;Caption table1.docx

Table 2

 2 Sex and age distribution of people interviewed in Ambendrana and Sahabe

Table

  Click here to access/download;Table;Caption table 2.doc

Table 3

 3 The social-ecological facets recorded in Ambendrana and Sahabe

Table

  Click here to access/download;Table;Caption table 3.docx

Acknowledgements

This study was funded by GIP Ecofor (Groupement d'Intérêt Public pour les écosystèmes forestiers). Funding to support the first author's stay in France was obtained from the Service Culturel de la Coopération Française of the French Embassy in Antananarivo. The Centre for Functional and Evolutionary Ecology (CEFE UMR 5175), CNRS in Montpellier hosted the first author and helped with data analysis and identification of Ficus species. We are grateful to the communities of Ambendrana and Sahabe for their willingness to participate in the interviews. We especially thank our guides Mr. Rajean Marie, Mr. Rakotomaro, Mr. Ramily and Mr. Roger for following us during our field studies and for accepting to participate in a workshop of the Intergovernmental Platform for Biodiversity and Ecosystem Services (IPBES) in Antananarivo (25-27 January 2016). We are grateful to Mr Dominique Hervé (Institut de Recherche pour le Développement) for local logistical support and to Mr Vonjison Rakotoarimanana for his support in data analysis. We heartfully thank Professor Doyle Mckey for his multiple comments on the manuscript and for having accepted to read the last version.

ECOFOR (Ecofor (Groupement d'Intérêt Public pour les écosystèmes forestiers).)