Local Knowledge of Utilisation and Threats Patterns of Termites around Pendjari Biosphere Reserve, West – Africa

Alain Kpèdékpo Gbeffe (  agbeffe@gmail.com ) Department of Environmental Management, Pan African University, Life and Earth Sciences Institute, University of Ibadan Thierry D. Houehanou Unversity of Parakou Francis A. Adesina Obafemi Awolowo University Musiliudeen A. Onilude Department of Agriculture and Environmental Engineering, Faculty of Technology, University of Ibadan Marcel B. R. Houinato Université d'Abomey-Calavi


Introduction
Biodiversity has fundamental values to human because of their dependence on it for nutritional, cultural, economic, and environmental/ecological well-being and its mismanagement leads to resource decline and biodiversity crisis [1]. Insects constitute a non-neglected component of biodiversity that deliver a host of ecosystem services fundamental to the survival of humankind [2]. Termites are social insects lling many ecological functions, especially in tropical ecosystems [3,4]. They play an important role in the decomposition of organic matter, soil fertilization, bioturbation and soil formation and vegetation growth and diversity in tropical regions [4][5][6][7]. They are important natural resources widely used in traditional medicine and also consumed by human populations in several parts of the world (Africa, Asia and America) [8]. Apart from being recognised as important pest and ecological engineer, termite species have therapeutic importance in traditional medicine worldwide [9].
In tropical and sub-tropical ecosystems, termite species are viewed as an important functional group for savannah structure and function, and their mounds have recently been shown to protect against drought by providing refuges for plants and foci for re-vegetation [10]. The bene ts and services associated to termites are however under-appreciated and more research is needed for a better capitalization of the importance of termite activity and their diversity in tropical ecosystems because ecosystems services provided by termites are not su ciently appreciated [5]. Given the importance of traditional knowledge in improving general understanding of insect ecology and biology, policies on the sustainable management and development of edible insect sector must take into account different roles that men and women in one hand, young, adult and old people in other hand play in edible insect management and involve them accordingly [2] for the sustainability of the bene ts and goods provided by those insects. Thus, Integrating local people's perception and knowledge in management strategies is important to guarantee local acceptance of the management plans of natural resources and biodiversity conservation [11] in one hand; and the sustainability of ecosystem services delivered by termite species particularly in tropical savannahs in another hand. Additionally, investigations on medicinal uses of animals (termites) and their products, as well as of inorganic materials, should not be neglected and should be considered as an important complementary body of traditional knowledge [12] and taken into account accordingly. Therefore, documenting ecological, uses knowledge and perceptions that local people have on termite species and their threatening factors is an important issue to be addressed. In this perspective a lot of works have been done and addressed the checklist of uses of termite species and their mounds [7,13,14] in savannah ecosystems without emphasizing on the pattern of those uses within the local communities for their better management. The other cultural signi cance and importance of termite species and their mounds were highlighted, apart from been foods, termite can also be used as feed for poultry or as bait to catch birds and sh [14]. The soldiers, the fungus gardens and the soil of termite mounds are also used for multiple medicinal purposes. Mounds and soil of termites have numerous functions: for geochemical prospecting, making bricks, plastering houses, making pots, and for storage. Termite mound soil is often used as fertilizer, agent for the rehabilitation of degraded lands, for restoring barren soil and for maintaining long-term soil productivity [13]. The mounds can also serve as burying places and are often associated with the spiritual world, especially containing the spirits of ancestors.
Termites also play a role as oracle, in superstitious beliefs, in art and literature. The main global impact of termites, outweighing their pest status, is clearly to provide the ecosystem services listed above [14]. From the ndings of all above-mentioned studies the question of knowing the pattern of knowledge uses of termites and their mounds is still relevant, due to the fact that studies that have examined the pattern of uses and the local people traits that affect uses and the perceived threats of termites within the local communities are very rare. The few conducted till now just made a checklist of uses regardless their pattern.
The present study therefore, aimed at analysing the use knowledge and threat factor patterns of termites and their mounds within the local communities around PBR. The speci c objectives associated to the study are (i) -to determine the diversity of termites and their mounds uses, (ii) -to assess the variations of range and evenness of uses knowledge among the local communities, (iii) -to examine sociolinguistic and demographic traits that govern local perception of anthropogenic activities, climate change and global use value on termite and termite mounds. In the case of medicinal plant species, age, gender, and dynamics of knowledge hypothesis, various individual socio-cultural and demographic traits such as gender, age, and literacy/formal educational level are all correlated with an individual's level of plant knowledge [15][16][17][18][19][20][21]. Age/age categories and gender are the most commonly examined variables [15] with women and older people tending to have greater knowledge of the local medicinal ora [15,21,22]. Thus, it had been suggested that (i) -Termite species and their mounds are well known and used differently by local communities around PBR, (ii) -women, older and indigenous people have greater and well distributed use knowledge of termite and termite mounds globally and as well as each use category compared to men, adult and young, and exogenous people respectively; and (iii)agricultural activities and construction uses are the main threats that in uence the sustainable uses of termite species and their mounds. The veri cation of those hypotheses will allow to know the key traits of the local community that should be considered in implementing eventual proposed management strategies of termite species and their mounds to sustain the associated good, services and bene ts that could maximize local community's well-being around PBR.

Study Area
The study was carried out in Benin, a West African country located between latitude 6°10'N and 12°25'N and longitude 0°45'E and 3°55'E, especially within the surrounded villages of Pendjari Biosphere Reserve (PBR). The Biosphere Reserve of Pendjari (PBR) located within Atacora district (10°30'-11°30'N, 0°50' -2°00'E) ( Figure 1) is one of the most important West Africa fauna reserves in term of plants and animals diversity conservation. It covers an area of 4800 km 2 and is composed of the National Park of Pendjari (2660 km 2 ), the hunting zone of Pendjari (1750 km 2 ), the hunting zone of Konkombri (251 km 2 ) and the controlled zone (139 km 2 ). The reserve was declared a Game Reserve in 1954, and upgraded to a National Park in 1961, and to a Biosphere Reserve in 1986 [23]. PBR is located in the Sudanian zone with one rainy season (April/May to October) and one dry season (November to March/April). The mean annual rainfall is 1000 mm with 60 % falling between July and September. The mean annual daily temperature is 27 °C. The vegetation is a mosaic of shrubs trees, woodland savannahs and grassland [24,25] regularly burned every year to provide fresh pasture to herbivores that dominate the reserve and increase visibility for touristic and hunting activities. Many small villages bound the Pendjari Biosphere Reserve. The population density in this area is low (38 inhabitants per km 2 ) compared with the whole country (87.2 habitants/km 2 ) [26]. Among the bordering local community of the PBR, Berba (65 %), Gourmantche (23 %) and Waama (7 %) are the dominant ethnics. The main activity of the local population is agriculture and animal herding. Women often exploit non timber forest products like fruits of shea trees among others, leaves of many species and also practice the rewood and charcoal trading.

Sampling of Informants and Data Collection
A pre-eld survey was conducted to determine the maximum sample size that was considered for the main survey. Thus 30 random respondents were selected in six randomly chosen villages among the villages surrounding PBR. Respondents were asked to provide answers to the following questions: "Do you know termite mounds and any services or goods provided by termites to human within the local community?" The proportion (P) of positive answers in each village was used to compute the corresponding sample size (n), while 10 % margin error (d) was considered. The sample size was calculated with the approximation of binomial law to the normal distribution [27].
Where N is the size of the sample and P is the proportion of informants who know and use the goods and services related to termites and termite mounds.
A total of 151 informants were chosen by using in each selected village the formulae of [27]. The different use types of termites and their mounds were recorded by using an individual questionnaire. The different categories of uses considered were foods or feeds, construction, medicinal, cultural and energy categories uses. Potential threat factors of natural resources and biodiversity particularly were previously identi ed from literature and respondents were asked to choose and rank them based on their knowledge and experience. They were asked to use the following de ned threat scale level: none (1), low (2), moderate (3) and high (4), to score each of threat factors which are agricultural activities, construction activities, vegetation re and climate change. The biodata (age, sex and the profession) as well as sociocultural data of the informants (ethnicity, level of study, and seniority in the village) were recorded. The informant's age was grouped into three categories young (15 ≤ age ≤ 30), Adult (31≤ age ≤ 60) and Old (age > 60). For the seniority, the con dence interval IC (29.95 ± 3.57) was built around the mean and used to de ne three classes of resident length (RL) within the community. Informants of low (RL ≤ 26 years), medium (26 < RL ≤ 34 years) and high (RL > 34 years) resident length were then de ned.

Data Processing and Statistical Analyses
For each reported use-type, the relative frequency of citation de ned as how often a use-report was mentioned was calculated using the delity level (FL %) [28]. The Fidelity level was assessed for both usetypes and use-categories. Where x is the number of informants who mentioned a speci c use or use-category and n is the total number of informants. The delity level was compared for different mentioned use-categories for both termites and their mounds using Chi square test.
Use knowledge of termite and termite mounds was measured using the relative use-value (UV) [29,30], a modi ed version of the use-value method introduced by [31]. This UV allows capturing all the known uses by an individual within and between use-categories [29,30].
Where UR uc,i is the number of use-report mentioned by informant i for a given use-category uc. In the dataset, UR uc,i varied from 0 to maximum (UR uc,i ) = 7, meaning that the maximum number of use-report mentioned by an informant in a use-category was 7. UV uc is the use-value for a given use-category uc which is the mean of UR uc,i for that use-category; n uc is the number of use-categories in the study (n uc = 5); n is the number of informants (151).
UV stands as a mean of URuc,i and could vary from 0 to n × n uc × 7 (in case all informants mentioned all use-categories and that all informants cited a number of use-report equals to 7 in each use-category). Because UR uc,i is a count data, a generalized linear model (GLM) with Poisson error distribution [32] was used to assess variation of UV (response variable) with respect to ethnicity, age category, and gender of informants (predictors). All predictors were categorical with respectively into four (Gourmantche, Berba, Waama and Others), three (Young, Adult and Old), and two (women, men) levels. Likelihood ratio test was used to assess the goodness of t of the nal model. The deviance-based pseudo-R 2 was also computed to assess the explanatory quality of the nal model.
The informants' diversity index and equitability value (table 1) were used to respectively assess the range and the evenness [33] of use knowledge of termites and termite mounds. In order to identify the key socio-economic factors that affect the use pattern of termites and termite mounds, GLM was implemented on informants' diversity index and equitability value related to termite and the termite mound uses. The indices vary from 0 to 1 [33,34], and their importance was revealed accordingly. Assess the evenness of knowledge [33] Source: [35] A pairwise ranking was implemented on the chosen threat factors to determine the factor that most in uence use knowledge of termites and termite mounds. The number of possible pairs was calculated using the formulae N (N-1)/2, where N is the number of chosen factors. Scores from each factor were summed up, ranks determined and the factors that received the highest total score were ranked rst.
All the statistical analysis was done by using the integrated development environment Rstudio (IDE) of R version 4.0.2 [36].

Socio-economic Characteristics of the Respondents
The   There was a highly signi cant (p < 0.01) relationship between overall knowledge of termite species and termite mound uses, and ethnicity and gender ( Table 4). The relationship is not signi cant between ethnicity and termite mound uses, whereas it is lightly (p = 0.052) between termite mound uses and gender.

Major threat factors of termites and termite mounds
Four major factors were selected for their signi cant threat on termites and termite mounds around Pendjari biosphere reserve according to local communities' perception. Agricultural activities characterised by slash and burning were ranked the top among the mentioned factors ( Figure 5) that affect termite, termite mounds and the associated good and services. They were followed in decreasing order by construction based on exploitation of dead termite mound, bush res which occur periodically each year and climate change as a global concern.
In uence of socio -linguistic and demographic traits on threat factors of termites and termite mounds   (Figure 6g) by humans especially the winged of Macrotermes genus or as feedings of poultry in general. This result is due to the fact that termites are widely consumed by people from all over the world because they provide the relevant amount of organic nutrients [37]. The utilization of Termites mainly Macrotermes as food, was also mentioned in some other tropical countries like Nigeria [38], Kenya [39], South Africa [40], and India [41]. In the villages surrounding PBR, the winged and roasted termites are the direct consumption form mentioned by informants. This was also observed in Nigeria [42] and in northern part of republic of Benin where winged termites are roasted and relished as a snack by people living the traditional lifestyle [4].
Termites especially Microtermes spp reported to be used as feeds for poultry ( Figure 6 j & k) in this study area, was not only reported in West Africa, but also in Burundi, Cameroon, Central African Republic, Congo, Madagascar and Tanzania [14]. The individuals of termite species, alone or in combination with their mounds in some cases, were also used to treat Umbilical ache, Fever, Bone fracture, and abscess in the study area which made medicinal use-category the second most cited use-category. These are in contrary with those of [4] who recorded no utilization of termites in traditional medicine within the adjacent phytodistricts to the present study area (Atacora phytodistrict). From the above-mentioned treated diseases using termites and the fact that medicinal uses of termites were also highlighted in many other part of the world, the ndings are supported by the evidence of antibiotic [43], antimicrobial [44,45], antifungal and antibacterial [46] properties of some termite species.
Apart from the direct uses of termite species, the respondents recognized the importance and cited some uses of termite mounds. The category of construction uses of termite mounds was the most cited one.
Thus, termite mounds especially the one of Macrotermes were used as construction or building material of furnaces, houses (Figure 6 e, f, h & i) storage bans pots, and housing for animal in general and poultry particularly. This remarks corroborated the one of Van Huis [14] who noticed that soil from deserted termite mounds is used for house construction in many parts of Africa like, in Togo termite mound soil is used to make furnaces in Akebu, Cotocoli, Ewe and Kabye communities. It is used as a kind of plaster for huts and for granaries to make the walls more resistant and smooth in Burkina Faso, Chad, Mali, Mozambique, Niger, Tanzania, Zambia and Zimbabwe. Likewise, the oor is done this way (Burkina Faso), and mixed with cow dung in order to make it really smooth (Mozambique, Sudan, Zambia, Zimbabwe). Medicinal and indirect food uses of termite mounds have also been mentioned by the local community of PBR. Thus, the intake of termite mound soil mixed with cow milk to treat the following health issues: the de ciency of iron in human blood, mumps, squirrel biting and umbilical dermatitis. The termite mounds are recognized as site for medicinal plant organs collection of some particular plant species (Figure 6 a, b), site for rewood collection (Figure 6c) and habitat for wildlife (Figure 6d).
Distribution of Use Knowledge pattern among local community and threats of termite and termite mounds around Pendjari biosphere Reserve Traditional knowledge analysis of termites, termite mounds and their global uses revealed that the information on use values of termites, termite mounds and their global uses were mainly in uenced by the sociolinguistic groups and the gender. Despite the fact this study is related to an insect (termites) nding aligned with gender, age and dynamic of knowledge hypothesis which suggests that various individual socio-cultural/sociolinguistic and demographic traits such as gender, age, and literacy/formal educational level are correlated with an individual's level of plant knowledge [15][16][17][18][19][20][21]. Concerning the gender, the ndings showed that Women are globally more knowledgeable of uses of termite, termite mounds and overall uses than men, and especially medicinal uses of termite, termite mounds. In many traditional societies, women are the primary healthcare providers in their households, so it is understandable their medicinal plant knowledge is greater than that of men [15], more they are the one in association with the kids in charge of collection edible part of plants and insects, among many other, for food purpose. These are in concordance with the nding of some previous studies on medicinal plant species [47,48] in general, and other multipurpose plant species such as Borassus aethiopum [30], Sclerocarya birrea [49], Tamarindus indica [50] and Adansonia digitata where women had more uses knowledge than men. This is in contradiction with the ndings on Cola nitida [35] where men had more uses knowledge than women. The authors justi ed by the fact that C. nitida is widely used for rituals often leaded by men traditional healer "vodounon". Those knowledge are well distributed among the two gender except termite uses where the knowledge were not well distributed among men. In another words the knowledge richness of uses of termite uses among men were detained by very few individuals.
For Ethnicity attributes, Gourmantche and Berba were more knowledgeable of termite and termite mounds uses due to the richness of their knowledge of medicinal and food uses of termite and termite mound in opposition to Waama and other sociolinguistic groups. This can be differently justi ed for Gourmantche and Berba sociolinguistic groups. Gourmantche rely only on natural resources and agro pastoral activities because they are enclosed between PBR and the Atacora mountain chain, and were inside the PBR before it was classi ed as game reserve in 1954, before it had been upgraded to Biosphere Reserve in 1986 [24]. More, they are located at almost 40 km from the main road "Route Inter Etat 2 (RIE2)" and don't have easy access to primary healthcare. This is in support with the ndings of Weckmuller et al [48] who stipulated that communities that are more physically isolated, inaccessible by road, and without easy access to primary health care, have a signi cantly higher use knowledge than those communities with a road and easy, free access to a Western doctor. The second group is the dominant sociolinguistic group of the region and located along the main road, thus they are in constant contact with other sociolinguistic group of the whole country and may gain a lot from this contact. The richness of knowledge on termites and termite mounds is similar for the considered age classes (young, adult and old), this is in contradiction with gender dynamic hypothesis of medicinal plant use knowledge which stipulated that older people have had much more experience with medicinal plant species over the course of their lives [51].
Threatening factors of termites and termite mounds around Pendjari Biosphere Reserve Agricultural activities have been pointed out as the factor that affect most termites and termite mounds for the sustainable uses of the associated bene ts, good and services. This was in concordance with the direct observations made on ground in one hand and to the ndings of some authors in other hands.
Firstly, agricultural activities in the study area are characterized by extensive slash and burning which require a lot of mass land to sustain an increasing population. This con rm the nding of Ackerman et al [52] that concluded termite species richness, abundance, and function often decline where forests are replaced by agricultural ecosystems (Agriculture, pasture and agroforestry systems). The agricultural activities are characterised by a heavy use of chemical mainly for the production of cotton in the area, and for yield improvement associated with the perception of pest status of termite species. This use of chemical is very detrimental to soil biodiversity in general and termites in particular when associated with the destruction of the termite mounds within agricultural lands because of the geographical position and status of the reserve don't allow the extension of the agricultural land by clearing natural vegetation [53] and was really stressed by riverine population during discussion.
Construction based on the use of termitaria soils because of its high clay content was also mentioned as major threat. This is the major construction material in so many villages around PBR. The promotion of construction of Voûte Nubienne (houses) based only on termite mound soils; which houses are more comfortable and adopted for the semi-arid and arid climates. Vegetation re is a tool generally use in the area for management purpose. It is used to clear vegetation around the buildings to avoid incidence due to accidental re and with the PA for hunting and tourism activities. Climate change in the other hands is a global concern and needs holistic actions to mitigate its effects on conservation of termites and their associated bene ts, good and services.
Ethnicity and age category are the two factors that in uence the perceived threats of agricultural activities and bush re on termite and termite mound. Berba are likely to perceive agricultural activities and bush re threats more than that of Gourmantche, Waama ethnics and others. Old informants followed by adults are likely to perceive bush re threats more than that of young's one. As threats of climate change are concerned, the perceived threats of climate change are in uenced by informant's age category and the reported global use values of termite and termite mounds. Thus, adults are likely to perceive climate change threats more than that of old and young informants in this order; and one unite increasing of termite and termite mounds' global use value, the informants' ability of perceiving climate change threats on termites and termite mounds increases of 17.6 % holding all others predictors constant. These ndings are in concordance with those of Norman et al. [54] who stated that age acts to increase an Individual's ability to perceive the threat. In another words, ability to perceive threats is accumulated along the life span because the perception is experience dependent [55]. It can also be justi ed by the fact that the extensive growth of agriculture is associated with increasing threats to biodiversity at all-time scales [56] including termites and all the associated goods and services.

Conclusion
The present study showed that the local people around PBR have different knowledge and use differently termites and (termite mounds). Termite species have food, feed and medicinal uses within the community, whereas their mounds have food, medicinal, construction, energy and cultural uses. Regardless the age, the sociolinguistic groups (ethnicity) and gender were identi ed as the traits that in uence the utilisation and use knowledge pattern of termites and their mounds around PBR. Gourmantche and Berba sociolinguistic groups were identi ed as the most knowledgeable of termite and termite mound uses, which knowledge is well distributed among them compared to the others, while women are more knowledgeable than men, and this knowledge is well distributed among women. Ethnicity and age category are the two factors that in uence the perceived threats of agricultural activities and bush re on termite and termite mound while the perceived threats of climate change are in uenced by informant's age category and the reported global use-values of termite and termite mounds.
Thus, in short terms, management action should be oriented towards the adults through sensitization on the importance of conserving termite species and their various habitats in order to make their associated bene ts, good and services more available for the people around PBR. Environmental education on the importance of termite species and their associated bene ts, good and services should be the key in the primary and secondary schools of the riverine villages around PBR. The promotion of trading roasted winged termites can also be an alternative diversi cation of household sources of income within the local community. Further studies are needed to analyse market value and the feasibility of trading roasted winged termites and access the proportion of living macrotermes mounds by including the ying distance of alates, the possible harvested quantity and their spatio-temporal variability. In long term, rstly implementing researches that can really de ne the quantity of termite mound soil that should be collected either on living or dead mounds in one hand and quantifying the real impact of each threatening factors on termites species biodiversity in relation to their importance in the ecosystems. Secondly, conceive a hand book (lexicon) on identi cation, recognition of PBR termite species and their uses in order to make those knowledge more available and transmissible from one generation to another.