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by Chris S. Duvall

A dissertation submitted in partial fulfillment of the requirements for the degree of

Doctor of Philosophy (Geography)


VILLAGES, VEGETATION, BEDROCK, AND CHIMPANZEES: HUMAN AND NON-HUMAN SOURCES OF ECOSYSTEM STRUCTURE IN SOUTHWESTERN MALI Chris S. Duvall Under the supervision of Professor Matthew D. Turner At the University of Wisconsin-Madison

This dissertation shows that both human activities and biophysical processes interact in complex ways to create an emergent ecosystem structure in southwestern Mali. This dissertation includes five body chapters. The first chapter is an analysis of settlement history in the research area, and situates the research in the context of current conservation practice in Malis Bafing Biosphere Reserve. This chapter shows that the indigenous Maninka people practice shifting settlement, and that frontier-style settlement expansion is not occurring in the area, as conservationists have assumed. The second chapter is an ethnographic study of Maninka physical geography terms, and shows that Maninka farmers perceive the landscape as highly heterogeneous, with few areas suitable for settlement or cultivation. The third chapter examines floristic patterns across the landscape, and shows that most floristic variation is due to edaphic features, especially the hydrogeology of a specific type of sandstone bedrock. Human activities have variable affects on vegetation, depending on various socioeconomic and biophysical factors. The fourth chapter shows that humans have affected the distribution of the baobab tree across the research area through activities that create suitable baobab habitat in settlement sites. The final body chapter shows that anthropogenic baobab groves represent important habitat for chimpanzees, and that conservation policies that affect settlement practice may reduce baobab regeneration and thus reduce chimpanzee habitat in the long term.

i Table of contents Acknowledgements........................................................................................................................ii Chapter One: Introduction .......................................................................................................... 1 Chapter Two: Settlement geography and biodiversity conservation in Malis Bafing Biosphere Reserve................................................................................................................. 12 Text box and figures for Chapter One .............................................................................. 49 Chapter Three: Folk taxonomy of physical geographic terms used by Maninka farmers in southwestern Mali................................................................................................................. 60 Figures for Chapter Two................................................................................................... 91 Chapter Four: Human and environmental causes of floristic patterns in southwestern Mali ............................................................................................................................................... 103 Figures, tables, and appendix for Chapter Three ............................................................ 149 Chapter Five: Human settlement and baobab distribution in southwestern Mali............. 183 Figures and tables for Chapter Four................................................................................ 211 Chapter Six: Chimpanzee diet, habitat use, and human settlement in Mali....................... 228 Figures and tables for Chapter Five ................................................................................ 257 Chapter Seven: Conclusion...................................................................................................... 272 Sources cited .............................................................................................................................. 283

ii Acknowledgements This research was completed with funding from the U.S. Fish and Wildlife Service Great Ape Conservation Fund, the Wildlife Conservation Society Research Fellowship Program, Conservation Internationals Primate Action Fund, the Zoological Society of Milwaukee County, and the Association of American Geographers Biogeography Specialty Group. Many people were crucial in my completing this dissertation. I especially appreciate the patience and hard work of my research assistants, friends, and hosts in Solo and elsewhere in southwestern Mali, who have endured my strange ways with constant hospitality and good humor. My advisor, Matt Turner, has provided several years of good advice, challenging questions, and generous amounts of time. All members of my dissertation committeeSara Hotchkiss, Lisa Naughton, Matt Turner, Tom Vale, and Karl Zimmererhave been generous with their time and have provided stimulating and valuable comments on my work. This dissertation is dedicated to my Grandma Elsie Tschetter, whose interest in everything about the world around her gave me the ability to complete my formal education as a student. All of my family, including ancestors I have not met, have contributed to my success as a student, and I wish to remember particularly my Aunt Aleta Bears and Grandpa Merle Duvall, who both inspired me and passed away while I was working on this dissertation project. All manner of support from my Mom, Dad, and brothers (and their families), as well as the thoughtfulness and generosity of my wifes parents, Earl and Susan Broidy, have been more important to me than I can express. Finally, my wife Jen and daughter Hazel have enabled me to complete this degree through their support and encouragement; my satisfaction in completing the degree remains far behind the satisfaction I get from spending time with them.

1 Chapter One: Introduction Human activities have profoundly shaped ecosystems in Africa. In the past century, humanitys long history in Africa and the ability of African farmers to sustain livelihoods in challenging environments were usually represented as causes for the continents purported environmental degradation. It is true that plant diversity in Africa is low relative to tropical Asia and South America (Richards 1973), but views about the human role in African ecosystems were, for much of the twentieth century, commingled with colonialist disrespect for African capabilities that hindered careful analysis of the environmental impacts of indigenous land management practices. Despite challenging environmental conditions, African farmers were not only portrayed as unsuccessful, but their purported ignorance of proper (i.e. European or North American) land management was seen as causing an ever-burgeoning degradation that made African environments so challenging (Bassett & Crummey 2003; Leach & Mearns 1996). This dissertation is an attempt to carefully assess the long-term impacts of settlement and farming on an ecosystem in southwestern Mali, and adds to a growing body of work that challenges the colonialist viewwhich has persisted in the post-colonial periodthat African land management is inherently destructive in semi-arid environments. However, the main contribution of this dissertation to human-environment geography is to show that a methodologically diverse approach is necessary to understand how human-environment interactions help create the emergent structure of ecosystems. The importance of this contribution is evident upon examining recent human-environment studies of long-term, anthropogenic environmental change in semi-arid Africa. These works have shown that the farming and settlement practices of smallholders often lead to vegetation changes that are not deforestationessentially the sole vegetation change recognized by earlier natural resource

2 scientists working in Africa (e.g. Aubrville 1949a; Schnell 1976; Stebbing 1938). Most prominent have been works attributing habitat enrichment to humans (Amanor 1994; Kreike 2003), especially the work of Fairhead and Leach (1996), who argue that farmers have created forest islands around settlements in savanna areas in Guinea, and that these forest patches were not the remnants of past deforestation as commonly believed by outsiders. Others have identified different vegetation responses to farming and settlementsuch as bush encroachment (Bassett & Boutrais 2000; Bassett & Koli Bi 2000), loss of species diversity (Devineau 2005; Lykke 1998; Nyerges 1989; Schreckenberg 1999), and resilience (Chidumayo 2004; McGregor 1994)but in nearly all cases these activities have not been shown to cause deforestation. Collectively, these studies represent the rise of an important new awareness in African geography of the diverse ways in which human societies have affected vegetation, and have helped chip away at the environmental degradation narrative that is deeply rooted in natural scientific discourse on Africa. Yet as studies of African ecosystems that include humans, these have all been limited in significant ways. In particular, this dissertation asserts that recent studies of the long-term, environmental impacts of settlement and farming in semi-arid Africa have had three specific, methodological problems that limit the accuracy and completeness of their analyses. This dissertation is built on a methodological foundation that reduces these limitations. First, several of these recent studies are built almost entirely upon qualitative analyses of environmental characteristics (e.g. Bassett & Boutrais 2000; Bassett & Crummey 2003; Fairhead & Leach 1996; Kreike 2003). The knowledge of local people, based on long-term observations of vegetation, is a valuable, and historically undervalued, source of information, and critical analysis of historic, scientific documents can identify persistent, inaccurate perceptions of

3 ecosystem characteristics held by outsiders. However, such qualitative sources are poorly suited to recognizing subtle biophysical processes, and to providing precise estimates of human impacts in spatial or ecological terms, thus limiting both the practical applicability and theoretical contribution of research findings. Furthermore, local knowledge, like scientific knowledge, is founded upon culturally specific perceptions and is not inherently more accurate than scientific knowledge (Agrawal 1995). Quantitative research methods can assess the accuracy of different perceptions of ecosystem conditionssuch as those of scientific and local knowledgeif these are designed in ways that do not privilege one set of perceptions over another. The specific methods used in this dissertationboth quantitative and qualitativeare described separately in each chapter in which a method is used. Broadly, however, quantitative methods are employed to test the assumptions of scientific viewpoints about specific aspects of the focal ecosystem, and the findings of quantitative analyses are interpreted using, and are used to interpret, local knowledge. This orderingtesting scientific knowledge then interpreting local knowledge was necessary because the local knowledge was not available when field research began, but its collection was an aspect of field research. Indeed, a second limitation of some recent studies of long-term, anthropogenic environmental change in semi-arid Africa has been an a paucity of local knowledge (Chidumayo 2004; Devineau 2005; Devineau 2001). Quantitative methods have certain advantages, but qualitative information gathered from local bodies of knowledge is necessary to situate ecosystem conditions in an appropriate sociocultural context. Socially and culturally determined beliefs, perceptions, and desires of land managers are primary determinants of the ways in which humans interact with other ecosystem components (Balick & Cox 1996; Croll & Parkin 1992). Local knowledge can provide rich detail on human environmental impactsif it is collected in a

4 manner that assures that it accurately represents the experiences and awareness of the focal population. An important way of doing this is to empower local people to identify, gather, and analyze information relevant to specific research questions. Much of this dissertation is based on data collection in which the residents of Solo village in southwestern Malisubsistence farmers representing the Maninka cultureactively participated. Although I write in a formal, third-person style throughout this dissertation, my personal history in Solo is important for understanding how and why the qualitative aspect of data collection and analysis developed as it did. From 1995 to 1997 I visited Solo several times as a Peace Corps Volunteer helping to develop the Bafing Biosphere Reserve, which was created in 1990, centuries after Solos founding. (A complete description of the research area is provided in Chapter 2.) Although I first visited Solo with a Malian forestry agent who enforced conservation policies with which Solos residents do not always agree, the c.250 people of Solo warmly received me, which encouraged me to return repeatedly on my own. These visits, along with experiences elsewhere in southwestern Mali, helped me develop friendships with several of Solos menmainly hunters with whom I share interests in wildlife, hiking, and natural historybased in part on my respect for their deep knowledge of the natural environment in the Bafing area. These relationships continued and improved in my subsequent visits to Solo, in 1999 (for my Masters research), 2002, and 2003, so that when I began field research for this dissertation in 2004, I had a decent understanding of how my proposed work fit into the body of concerns held by people in Solo. In particular, Solos people are concerned about the security of their access to land and other resources, which is threatened by continued development of the Bafing Biosphere Reserve. I have also maintained professional relationships within the Malian conservation community.

5 Having a window into the worlds of these conservationists and the people in Solo (and nearby villages) has provided a curious view of human-environment interactions. The conservationists have generally seen the land management practices of people in Solo as destructive of biodiversity resources, while people in Solo see their activities as part of a strategy implemented over generations to sustain the productivity of diverse domesticated and natural resources. These viewpoints have meant that the conservationists have sought to modify or prohibit various indigenous practicesto attain their goal of stopping purported biodiversity losswhile people in Solo have sought to maintain indigenous practices while avoiding fines and other negative consequences of conservation law enforcementto attain their goal of sustained productivity of diverse resources. Before beginning this dissertation, I felt that neither group had a complete understanding of the processes that helped create the biodiversity upon which both were focused, although I believed that people in Solo certainly had a more complete knowledge than city-based conservation bureaucrats. Thus, when I began field research in 2004, I explained to both groups that I sought better knowledge of the processes that created biodiversity across the Bafing landscape, and particularly how the activities of people like those in Solo affected these processes. Predictably, both groups were certain that I would confirm their views, and many people in these groups willingly helped me collect data I asked for or they thought I should have. Since I lived in Solo, though, I gave the people there much more opportunity to contributeI certainly felt that they had the most to share. Furthermore, since conservationist perspectives have dominated resource management policy in the Bafing reserve (i.e. Caspary et al. 1998; PREMA 1996; i.e. Warshall 1989), I felt that it was important to gain a different perspective on resource use, that of the people in Solo. In 2004 I lived in a thatched hut in Solo that had been first offered to me as a

6 visitor in 1995, shared meals and experiences with my host and his family, and spent my time working and socializing with friends and neighbors. Solos people participated in my research in three main ways. First, many people willingly responded to my questions (which were, to them, often tedious and nave) about plants, animals, local history, and other topics, and several people volunteered additional information I had not specifically asked for, but which they felt was important for understanding the people-vegetation-wildlife relationships of interest to me. Second, Solos traditional authoritiesthe chief and several men serving as counselorshelped me identify and map all settlement sites, permanent water sources, and forest patches by volunteering information far more specific than I could have asked. Third, and perhaps most importantly, six men I hired as research assistants undertook the difficult task of searching large portions of the landscape for chimpanzees twice per week. Their knowledge of the landscape and of chimpanzee behavior allowed me to amass significant amounts of data on chimpanzees. Overall, the participation of Solos people in this research was crucial for its completion, as well as the completeness of its content. The focus on chimpanzees in this dissertation suggests a third limitation of recent humanenvironment studies of the impacts of smallholder farming and settlement on semi-arid African ecosystems. None of these studies have substantially considered impacts on trophic levels other than vegetation. Of course, the effects of drastic, short-term vegetation changes on wildlife have received attention because of the rate at which commercial logging is transforming African rainforest environments (Johns 1982; Johns & Skorupa 1987; Plumptre 2001; Plumptre & Reynolds 1994; Skorupa 1986). However, the effects of subtle, long-term vegetation changes on wildlife in Africa are only generally known (Happold 1995), and only a handful of papers have addressed this issue directly, in rainforest environments (Fimbel 1994b; Wilkie & Finn 1990).

7 Throughout the semi-arid tropics, very little attention has been given the effects of subtle, longterm vegetation changes on wildlife (cf. Bourlire 1983; Bullock et al. 1995; Cole 1986). If human-environment geographers accept that many natural landscapes are pervasively humanized (Zimmerer & Young 1998), then greater attention must be paid to identifying and understanding anthropogenic features of animal, and not just plant, communities (cf. NaughtonTreves 2002). The spatial structure of an ecosystem results from interactions between its biotic and physical components. Humans dominate many ecosystems, but in all ecosystems processes that operate independently of humans constrain interactions between biotic and physical components (Vale 1982; Zimmerer & Young 1998). Thus, possible human impacts on ecosystem characteristics are limited, even if such limits may have little meaning in urban areas. In rural, agrarian landscapes, though, biophysical limits on the range of possible human environmental impacts are more important, and their identification is crucial to determining how human activities contribute to ecosystem structure and function. The difficulty of identifying these limits is perhaps the most important reason why mixed methods should be used in humanenvironment geography. Indeed, by substantially analyzing biophysical as well as sociocultural components of ecosystems and environmental change, human-environment geographers can also advance biogeography. Only recently have scholars begun to study biogeographic effects and processes of anthropogenic vegetation change in Africa using sophisticated sociocultural evidence to support biogeographic arguments (e.g. Assogbadjo et al. 2006; Maranz & Wiesman 2003; O'Brien & Peters 1998). This dissertation advances both human-environment geography and biogeography by using mixed methods to address significant questions on how human activities contribute to ecosystem structure. It is not simply a cultural ecology informed by

8 biogeography, but an analysis of the biogeography of human activities in a landscape where the ecology and geography of these activities are poorly known. The body of this dissertation is written as five related, but independent, papers, and not as a single text divided into chapters. Each paper develops unique arguments and themes, but several themessummarized in the following paragraphsrun across several chapters. Each paper contributes to the overall argument of the dissertation, that settlement and agriculture have increased the distribution and abundance of chimpanzee habitat in the focal landscape. Figures and tables associated with each chapter are placed at the end of chapters, but a single bibliography is provided at the end of the dissertation. The narrative this dissertation traces is about how Malian society, Maninka culture, and the biophysical environmental have contributed to the spatial structure of an ecosystem in southwestern Mali. Its chapters range from a broad analysis of settlement history and conservation practice to narrowly focused analyses of specific plant and animal components of the research area, but all chapters are fundamentally about the spatiality of the patterns and processes observedwhere things are and happen across the landscape. The following chapter provides a detailed description of the research area and of the conservation policy context in which the research is situated. By providing an analysis of settlement history and practice in Malis Bafing Biosphere Reserve, along with an outline of ongoing conservation policies meant to address the threat settlement is supposed to represent, this chapter exposes the tension that exists between local and scientific knowledge of anthropogenic environmental change. This tension is a primary element in the dissertations overall narrative. The main argument of Chapter 2 is that conservationists have misunderstood the threat to biodiversity posed by indigenous settlement practices because they have assumed incorrectly that the spatial pattern of

9 settlement indicates a process of frontier-style population expansion. This process has not occurred for centuries in this landscape, and policies based on the assumption of ongoing expansion are likely to fail conservation goals in the long term. Chapter 3 narrows the focus of the narrative by providing an ethnographic analysis of Maninka physical geographic concepts. This chapter provides an important contrast with technical, scientific concepts in physical geography, which are more familiar to the reader and the topic of Chapter 4. Chapter 3 shows that Solos residents do not perceive biophysical diversity across the landscape in the same way as natural resource scientists. Maninka farmers perceive detailed variation in the abundance and accessibility of resources across the landscape based not only on tangible characteristics, but also on intangible, socially determined characteristics. Based on this perceived variation, different parts of the landscape are subject to different use. Maninka farmers do not consider most portions of the landscape arable or suitable for settlement. This chapter contributes to the dissertations overall narrative by showing that the conceptual landscape upon which Maninka settlement practices are founded does not include any type of frontier along which human activities are increasing. Instead, the Maninka conceptual landscape is a mosaic of areas with different histories of use, and varying potentials for use. In Chapter 4, the overall narrative moves through a quantitative assessment of the scientific view that human disturbance is the major cause of floristic variation across the Bafing landscape. Using floristic analyses of extensive vegetation samples in sites with known disturbance history, this chapter identifies biophysical and human factors that are significantly associated with variation in vegetation composition. This chapter underscores the significance of edaphic features as the primary source of floristic variation. Edaphic features are, of course, fundamentally important in plant biogeography, but in the West African context their importance

10 is frequently ignored or underemphasized because landscapes are often portrayed as profoundly humanized. The physical structure of the sandstone that outcrops across the research area creates highly distinctive plant habitats that are associated with elevated biodiversity. Settlement and cultivation also affect vegetation composition, but only in sites with relatively deep, arable soil, and not across the entire landscape. Additionally, in sites with deep, arable soil, the effects of settlement and cultivation are variable, ranging from species enrichment to species decline. Chapter 4 shows that human activities have altered the distribution of various economically important tree species, especially those with edible fruits; Chapter 5 tests this possibility for a single species, the baobab (Adansonia digitata). This chapter is based on point-pattern analysis of a census of all baobabs and settlement sites in the research area, and tests two different scientific views on the reason for apparent spatial correlation in the distribution of settlements and baobabs across Africa. Chapter 5 shows that human activities lead directly and indirectly to the creation of baobab groves at settlement sites. The overall narrative is maintained in Chapters 4 and 5 as a study of how the direction and intensity of anthropogenic effects on vegetation characteristics can be precisely determined, in order to understand how scientific and local perceptions relate to observed patterns across the landscape. Both chapters build explicitly on the findings of Chapters 2 and 3. Finally, in Chapter 6 the overall narrative refocuses on the tension that exists between local and scientific views of human environmental impacts, and continues the theme explored in Chapters 4 and 5, on how to precisely determine anthropogenic effects on ecosystem characteristics. Specifically, Chapter 6 examines chimpanzee distribution and behavior in relation to human settlement history (Chapter 1) and vegetation characteristics (Chapters 4 and 5) in the research area. The quantitative and qualitative analyses in Chapter 6 show that

11 chimpanzees frequently visit baobab groves at abandoned settlements during the time of year when baobab fruit composes an important component of their diet. However, through most of the year, habitats along sandstone outcrops are more important habitat, due to the abundance of food plants and surface water in these areas. Baobab groves at settlement sites are most frequently used when food abundance in cliff habitats is lowest. Thus, human activities expand the distribution and abundance of chimpanzee habitat relative to that which exists due to biophysical processes operating independently of humans. The narrative of this dissertation finishes with a short conclusion that returns to the conservation context described in Chapter 2. The concluding Chapter 7 underscores the importance of identifying and understanding the biophysical and sociocultural context in which human-environment interactions occur. A primary argument of this dissertation is that there has been insufficient attention given to the spatial and temporal contexts of indigenous land management in the research area in particular, and elsewhere more generally. Better use of mixed research methods in human-environment geography is necessary to improve knowledge of the contexts of human-environment interactions. By recognizing how context constrains both the environmental impacts of human activities and the appropriateness of possible conservation interventions, human-environment geographers can improve the long-term effectiveness of biodiversity conservation practice.

12 Chapter Two: Settlement geography and biodiversity conservation in Malis Bafing Biosphere Reserve

Abstract Studies of rural settlementdefined formally as a distinct land use dedicated to human shelterhave been more important in cultural than in human-environment geography, although settlement practice is constrained by the socioeconomic and biophysical processes that dictate natural resource use. This paper argues that recognizing settlement as a distinct land use improves our ability to understand and manage human environmental impacts. An analysis of settlement history and Maninka settlement practice in part of Malis Bafing Biosphere Reserve shows that the Maninka practice a shifting, as opposed to fixed, settlement system that allows economically and politically marginalized men to improve their access to natural resources without threatening traditional political authorities. However, conservationists have viewed Maninka settlements as fixed, rather than shifting, so that settlement pattern has been misinterpreted to suggest a process of frontier-style settlement expansion, and associated wildlife habitat loss. This viewpoint is inaccurate, and has led to conservation policies that create hardship for the marginalized people who benefit most from shifting settlement, and neglect the geography of biodiversity and settlement in the Bafing area. These policies will likely fail conservation goals in the long term. Keywords: settlement; conservation; Mali; Bafing; chimpanzees

13 Introduction Cultural geographers working have long studied the form and distribution of rural settlements as a means of understanding perceptions and use of space, demography, historical processes of landscape change, and spatial aspects of agricultural and economic development. In the West African context, settlement geography has been studied primarily as a means of understanding cultural geography and history (e.g. Bernus 1956; de Bruijn & van Dijk 1995; Gado 1980; Gallais 1975; Queant & de Rouville 1969; Sidikou 1974; Woodford 1974), or constraints on economic development (Silberfein 1998). Human-environment geographers have given less attention to settlement, even though settlements are established and abandoned due to socioeconomic and biophysical processes that affect resource use (Stone 1996). Instead, humanenvironment geographers have focused on more directly productive types of rural land use such as animal husbandry, agriculture, logging, and conservationwithout specifically examining settlement practices associated with these other land uses. Recently, a handful of human-environment anthropologists have published valuable studies of settlement ecology (Amanor 1994; Fairhead & Leach 1996; Stone 1996) that build on important, earlier works in anthropology (de Schlippe 1956; Green et al. 1978). In geography, however, there have been few published analyses of rural settlement from a human-environment perspective since about 1970, the major exception being Chisholms (1979) third edition of Rural Settlement and Land Use, originally published in 1962. From a geographical perspective, our knowledge of rural cultural and political ecology is built largely upon studies of land uses other than settlement. Settlementformally defined as the development of human shelter over time through various social, cultural, spatial, and ecological processes (Chisholm 1979; Christaller 1966 [1933]; Hill 2003; Stone 1996)is a distinct land use (Morgan 1955). Of course, settlement is

14 not entirely separable from other land uses, especially agriculture, husbandry, logging, and reserving land for hunting and gathering, or recreation. However, settlement practice includes many acts that are not part of other types of land use, because shelter is the primary purpose of settlement, and no other land use. For instance, in settlements, people must identify, construct, and manage places for resting, cooking, storage, caring for children and the elderly, bathing, gardening, keeping small animals, and many other activities that are less frequently, if ever, practiced in places dedicated to other land uses (e.g. Morgan 1955). Settlement is ecologically the most disruptive land use in agrarian rural landscapes, though it directly affects small areas; settlement sites are occupied more continuously and are more intensively managed than other parts of the landscape (Stone 1996). More importantly, settlement constrains the spatial distribution and intensity of most other land uses (Stone 1996). Farmers seek settlement locations that minimize the distance between settlements and fields, fallows, water sources, construction materials, and other resources (Chisholm 1979; Hill 1953; Jarrett 1948; Johnson 1977; Morgan 1955). As a result, resources near settlement sites experience heavier use than more distant ones (Stone 1996). Recognizing and studying settlement as a distinct land use requires spatial and temporal scales of observation that are infrequently used in human-environment geography (Stone 1996). Cultural ecologists often focus on changes that occur over periods up to about a decade, the maximum being about the length of time an individual swidden remains productively fertile. In contrast, the periodicity of settlement establishment and abandonment in non-irrigated, agrarian landscapes is generally decades to centuries (Hill 2003; Hunter 1967; Udo 1965). Of course, proxy data have allowed many cultural ecologists to use very long periods of observation, but such observations generally occur at regional or continental scales that obscure events and

15 processes occurring in specific communities or landscapes. Community-based cultural ecologies of agriculture often focus on questions at the spatial scale of individual plots or settlements: What prompts farmers to abandon one plot and move to another? What leads to intensification in one plot or village but not another? However, settlement occurs at a more expansive scale, across landscapes measured in tens to hundreds of kilometers that include areas that are suitable for settlement, but not occupied (Chisholm 1979; Hill 2003; Hunter 1967; Morgan & Woods 1986; Stone 1996). The scale of observation suitable for studies of settlement ecology is more similar to that often used in studies of pastoralism (e.g. Bassett 1988; Bassett & Koli Bi 2000; Behnke et al. 1993; Coppolillo 2000; Turner & Hiernaux 2002). The present paper has two goals. Its primary goal is to describe the settlement practices of Maninka farmers in southwestern Mali and argue that these subsistence agriculturalists practice a system of shifting settlement (Stone 1996). This system is an adaptation to a particular set of socioeconomic and biophysical conditions that are widespread in Africa, where shifting settlement appears to be widely practiced (de Schlippe 1956; Hill 1953; Hunter 1967; Netting 1993; Richards 1978; Stone 1996). This description of Maninka settlement practice is important because shifting settlement is ethnographically underdocumented (Stone 1996), although it is widespread in space and time (Murdock 1967). Cultural geographers of settlement appear to have also largely overlooked it (cf. Grover 1985; Hill 2003; Kharkwal & Sharma 1990; Sharma 1985) even though Stone (1996) shows clearly how shifting settlement systems may arise based on models of agricultural, geographic, and demographic change that geographers widely use. In short, building on Chisholms (1979) model of settlement location and Boserups (1965) model of agricultural intensification as modified by Brookfield (1972; 1984), Stone (1996) argues that shifting settlement systems develop where the following factors co-occur: a) farmers face a

16 shortage of farmland within a reasonable distance of their settlement; b) unused, but relatively small, patches of farmland exist some distance away from their settlement; and c) the costs of agricultural intensification in fields near their settlement exceed the costs of abandoning the settlement and building a new one near an unused patch of farmland. Farmers in shifting settlement systems probably always practice shifting cultivation (cf. Morgan 1955), but these two types of land use are distinct and occur over different spatial and temporal scales. Maninka shifting settlement complements Maninka shifting agriculture and allows farmers to adapt to socioeconomic changes in a patchy biophysical environment. However, policy makers have not recognized that the Maninka practice shifting, and not fixed, settlement, and thus have misinterpreted Maninka settlement pattern as an indication that frontier-style settlement expansion is occurring (Caspary et al. 1998; PREMA 1996). Conservationists view the relatively numerous, new settlements in Malis Bafing Biosphere Reserve (BBR) as the disintegration of larger settlements as people move into previously wild and unoccupied land. Thus, settlement is seen as an unqualified threat to wildlife habitat. The second goal of this paper is to challenge this viewpoint by examining the history and practice of settlement in a portion of the BBR. This examination shows that: a) most settlements are shortlived; b) virtually all settlements have been established in sites that previously hosted now abandoned settlements; and c) shifting settlement is a flexible institution that enables Maninka farmers to adapt to socioeconomic and biophysical change in an environment with sparse, patchy resources. The paper concludes that understanding the geography and rationality of shifting settlement will enable conservationists to manage biodiversity resources more effectively while also reducing the social injustice, and probable long-term failure, of current policies meant to reduce wildlife habitat loss to human settlement.

17 Geographical Context Research occurred in an area of 183 km2 around Solo, a settlement of about 200 people located in the Bafing Biosphere Reserve (BBR) of southwestern Mali (Figure 1, p. 52). The BBR comprises two national parks and a chimpanzee-specific reserve; a buffer zone surrounding these areas is awaiting ministerial approval (Duvall et al. 2003). Solo lies on the northern boundary of one of the national parks, so that about half of its traditional territory lies in the park, while the remainder lies in the proposed buffer zone (Figure 2, p. 53). The BBR protects an important population of West African chimpanzees (Pan troglodytes verus) (Duvall et al. 2003), as well as populations of several threatened or endangered trees (Duvall 2001). Chimpanzees have been a focal species for conservation activities in the BBR since the mid-1980s (Maldaque 1985; Moore 1985), and this focus will certainly intensify with increasing international recognition of the significance of Malis population (Kormos & Boesch 2003; Kormos et al. 2003). The plant and animal species of greatest conservation concern are associated with sandstone outcrops that rise 200-300 m above surrounding, relatively flat lowlands (Duvall 2001; Duvall 2000; Jaeger 1959; Lawesson 1995). Topographic complexity in these outcrops helps create a wide range of microhabitats, and the physical structure of the rock creates very stable ecological conditions (cf. Larson et al. 2000). Erosion of the sandstone plateaus has formed narrow ravines, rocky slopes, and plains with relatively infertile sandy and silty soils. The upper surface of the plateaus consists of bare or shallowly buried bedrock (Jaeger 1950b; Jaeger & Jarovoy 1952), and similarly xeric ferricrete hardpans are common throughout the area (Dames & Moore 1992; Michel 1973). The best farmland is located in small, basin-shaped valleys below the outcrops (PIRT 1983; Samak et al. 1987). Permanent springs are most common along the

18 sandstone outcrops, where sedimentary layers in the sandstone have been exposed (see Chapter 6). Elsewhere, permanent surface water sources are uncommon, and are primarily deep depressions in seasonal streambeds. People rely mainly on hand-dug wells for their water needs. Precipitation is highly seasonal and averages about 1100 mm per year, with high interannual variation (FAO 1984; Leroux 2001; PREMA 1996). Woodland vegetation dominates most of this landscape, especially in areas with relatively deep, fertile soil. Forest patches occur in topographically protected microhabitats with moist soil conditions along the sandstone outcrops. Locations with shallow or infertile soil host patches of edaphic bushland or grassland. Based on woody species composition, fifteen vegetation types have been described for the area, including several types that are associated primarily or uniquely with abandoned settlement or field sites (see Chapter 4). Although gazetted originally in 1990, the BBR remains almost non-existent on the ground. The Malian government, supported by bilateral aid agencies, has developed and adopted a formal management plan for the area (Caspary et al. 1998; Niar 2000). The management plan identifies Maninka settlement practices as one of the main threats to biodiversity in the BBR because it causes wildlife habitat loss (Caspary 1999; Niar 2000; PREMA 1996). In particular, Maninka farmers establish hameaux de culture (farming hamlets) away from their official villages of residence in order to access patches of arable soil (Ciss 1970; Koenig & Diarra 1998; Samak et al. 1987). For clarity, the French terms hameau and village will be used when referring to a hamlet or village from the administrative viewpoint of Malian conservationists and government officials, to distinguish these concepts from the Maninka terms bugu [hamlet, farm] and dugu [village], described below. The English terms hamlet and village are used interchangeably with bugu and dugu. Hameaux are generally small, occupied by just a few

19 nuclear families, although some are as large as a village, hosting many families (PREMA 1996). Most distressing to conservationists is that many hameaux have been established throughout the BBR in the past ten years, and that some hameaux, like villages, have been occupied for decades (Caspary et al. 1998; Niar 2000; PREMA 1996). Hameaux are not officially recognized settlements; their residents are counted as part of the population of the villages where they lived before moving to a hameau. The only legal or administrative status most hameaux have is as illegal settlements, if located in the BBR, or illegal clearings, since Malian forestry laws prohibiting clearing vegetation that has not been cleared for 10 years (Prsidence de la Rpublique du Mali 1995). Conservationists working in the BBR consider hameaux de culture a spatially uniform cause of habitat loss that occurs in an essentially chaotic manner through most of the reserve (Niar 2000; PREMA 1996). Indigenous land use is believed to happen without any planning beforehand, and the establishment of new hameaux is considered an uncontrolled swarming (1998: 98). No study of settlement practice has been made in the Bafing area; the information used to support this viewpoint is primarily PREMAs (1996: 45) list of 83 hameaux associated with 8 villages in the BBR area, which also shows that many of these have been occupied less than ten years. Few of these hameaux have been mapped, although PREMA reports that many are >5 km from their associated villages. This distance is significant, because the presidential decree that originally established the BBR (as a game reserve) in 1990 acknowledged the existence of some, but not all, preexisting villages in the protected area, and allowed these villages circular enclaves 5 km in diameter, in which farming was allowed (Prsidence de la Rpublique du Mali 1990). Several preexisting villages were not recognized, and some of the villages that were recognized are

20 actually hamlets (i.e. Maninka bugu), apparently because those who drafted the proclamation had very little information about local geography. The existence of other hamlets in the BBR was not recognized. In fact, all settlements in the BBR not in the recognized enclaves became illegal with the reserves creation, even though many occupied in 1990 had been occupied for decades. From the beginning, human presence in the BBR area was erroneously underestimated and spatially simplified: lawmakers envisioned that existing human settlement could be encapsulated in a small number of standardized, circular enclaves. Greater familiarity with the Bafing landscape, which came as a result of various development projects in the 1990s, caused conservationists to discover that there were many more settlements in the BBR than previously imagined (Duvall & Niagat 1997; PREMA 1996). The increased observation of settlements was interpreted as [t]he populations geographic expansion (Caspary et al. 1998: 84), even though there was no relevant information reported on the historical geography of settlement or on demographic change for the area. For instance, population estimates for the BBR area, based on rapid surveys in many settlements, are paired with population growth rates for western Mali or all of Mali (stated as 2-4% annually) to suggest that the population in the BBR is increasing at the same, relatively high rate (Caspary et al. 1998; PREMA 1996). This method of analysis obviously combines two different scales of information. More careful demographic analysis has shown that the human population has declined in the Bafing area since the mid-1980s, and may continue in this direction for the next several decades (Mission Francaise et al. 1996; Raynaut 1997), even though regional population centers are growing (Bonavita 2000). Nonetheless, the expanding population interpretation of the observed settlement pattern led conservationists to recommend revising the enclave strategy to address this perceived

21 problem of habitat loss. Residents would be allowed to farm within traditional territoires des villages (village territories), which were to be delimited in collaboration with residents, but outside these areas the level of resource protection would increase to make integral sanctuary zones for wildlife, as defined by IUCN (1986), in which no activities (other than tourism and research) would be allowed (Caspary et al. 1998: 84-86). The likely size of village territories was not specified (although they were suggested as replacements for the 5-km enclaves), and the acceptability of hameaux within village territories was not addressed. Although no village territories have been identified, the integral sanctuary zones were created in 2002, when the two national parks, Kouroufing and Wongo, were gazetted (Prsidence de la Rpublique du Mali 2002a; Prsidence de la Rpublique du Mali 2002b). These protected areas have become nationally significant in Malis efforts to meet its biodiversity conservation goals. Under the Convention on Biological Diversity, the Malian government initiated the Project for the Long-term Management of Biodiversity in the Bafing [Biosphere] Reserve (MEATEU 2000: 83-84). One outcome of this project was the governments eviction, in 2004-06, of all hameaux in the BBRs two national parks, affecting an unknown number of people. Threats of violence were used to evict settlements: in January 2006, agents of the national conservation directorate threatened publicly to burn any hameaux remaining in the BBR. Evictees have received no compensation or assistance. Most have returned to their home villages, or established new settlements just outside national park boundaries. Conservationists continue to fear the settlement expansion hameaux are supposed to represent, and people living in settlements in and near the BBR fear the possibility of more evictions in the future.

22 Settlement geography in the BBR has not been studied, despite the certainty conservationists have expressed in representing the relationship between settlement pattern and process. An examination of settlement history and practice in the BBR shows that there has been no geographic expansion of settlement for centuries. Settlement pattern represents an ongoing process of settlement establishment and abandonment observable over decades, not years, arising from a social institution that enables politically and economically marginal families greater access to natural resources without threatening more dominant families. Data collection and analysis Data on settlement history and practice were collected through documentary sources, ethnographic interviews and participant observation, and foot surveys of abandoned settlements in the research area. Documentary evidence provides some information on settlement history in the research area. The Scottish explorer Mungo Park passed through Solo during his second visit to what is now Mali, in 1805 (Park 1954 [1815]). Other European travelersmainly French military officersvisited other parts of southwestern Mali in the 1800s, recording observations relevant to understanding general settlement history in the area (Mage 1868; Mollien 1820 [1967]; Noirot 1885; Tellier 1898). During the 1880s-1890s, the French gained military control over western Mali, and in 1889 a colonial officer collected names of occupied settlements in part of the research area and to its northeast, along the Bafing River (Samak et al. 1986). In 1952, the French colonial mapping service took aerial photographs of southwestern Mali, and produced 1:200,000 scale topographic maps from these images (Anonymous 1958). While many settlements are unnamed on this map, the map and photographs record the existence of many settlements in 1952. Finally, rapid surveys of settlement history around Solo were conducted in

23 the 1980s as part of the resettlement projects associated with construction of the Manantali Dam (Samak et al. 1986; Sanogo 1991). Oral history has retained much more information about past settlement around Solo. Oral historical interviews were conducted in Solo, in the Maninka language, during May-July 2003 and January-December 2004. These interviews built upon the researchers past visits to Solo since 1995, including visits to now-abandoned hamlets. Abandoned settlements were identified during individual interviews with c.45 male and female residents of Solo aged c.10-80, and during three group interviews of Solos traditional authorities (chief, land chief, and senior counselors), all men aged c.40-80 years. With the assistance of interviewees, all occupied and abandoned settlement sites identified through interviews were visited. The location of each site was determined using a Garmin GPS-12XL unit, and was recorded as a point corresponding to the approximate center of the occupied area of each site (as evidenced by the distribution of huts or remains of hut foundations). For every site, the following information was collected: site name, estimated dates of establishment and abandonment, causes for establishment and abandonment, soil texture (via manual analysis: Midwest Geosciences Group 2003), and preceding and succeeding residences of site occupants. Multiple informants were interviewed to increase precision by triangulating date estimates and gaining multiple perspectives on other information (cf. Flowerdew & Martin 1997). Generally, establishment and abandonment dates were estimated by correlating informant life history markers, changes in site occupation status, and datable events, such as national elections. In some cases, specific dates of past site occupation were gathered from the historic documents and aerial photos described above. The resulting point pattern was analyzed using Ripleys univariate K function. There are numerous technical descriptions of Ripleys K, which is calculated from the number of points in

24 a given distance, h (e.g. Bailey & Gatrell 1995; Diggle 2003; Dixon 2002; Haase 1995). In application, the K function is usually linearized to stabilize variance and facilitate interpretation, and in this form is called the L function (Dixon 2002). Additionally, multiple values for h are used in order to assess spatial pattern at multiple scales. Values of L(h)>0 indicate clustering, while L(h)<0 indicates regularity in the distribution of points (Dixon 2002). Since the present application of Ripleys K is solely to describe the spatial structure of the observed point pattern and not to assess statistical significance, the assumption of stationaritywhich is probably not held for the distribution of settlementsdoes not need to be met (Bailey & Gatrell 1995; Diggle 2003). K function analyses were conducted using the SPLANCS package (version 2.01) in the R statistical software environment (version 2.2.1). For further description of SPLANCS, see Rowlingson and Diggle (1993), Gatrell et al. (1996), Bivand and Gebhardt (2000), and Diggle (2003). Edge effects were corrected geometrically (Bailey & Gatrell 1995). Finally, ethnographic interviews and participant observation were used to identify and understand Maninka settlement practices. Participant observation provided experiential knowledge of settlement practices, while interviews clarified observations (Werner & Schoepfle 1987). During the period of research, four settlements were abandoned (three as a result of eviction from the BBR) and one was established (outside the national park by residents of an evicted hameau). Informal interviews were conducted while working or socializing with farmers, and while working with men who were clearing the site of the newly established settlement. All but two of c.45 intervieweesmales and females aged c.10-80had previously lived in one or more now-abandoned settlements, and all were subsistence farmers. Settlement history before 1890

25 Current Maninka settlement practice developed during the settlement history of the research area, which can be divided into three broad periods: prior to 1890, 1890-1960, and 1960 to the present. Within these broad periods there are several distinct phases, which are described individually in the following paragraphs. Early Holocene. During the early Holocene and before, modern humans settled the area around Solo, as evidenced by stone artifacts and cave shelters (Sanogo 1991). Essentially nothing is known of these people or their settlement practices. Pre-Maninka period. At an unknown period, the area was settled by people the Maninka identify as Jan, Sylla, Sig, and Somalaka (Samak et al. 1986). The Maninka entered the research area at an imprecisely known date following the Manding expansion in Sudanian West Africa, which peaked in the 1300s (Ciss 1970; Samak et al. 1986). According to local oral history, the Maninka militarily defeated the earlier residents, causing the earliest cases of settlement abandonment for which there is historical evidence (Figures 2 & 3, pp. 53 & 54). Early Maninka period. The Maninka have never occupied the settlement sites from which they expelled the Jan, Sylla, Sig, and Somalaka, some of whom fled, while others were absorbed into Maninka society as hereditary blacksmiths, leather workers, and griots (Samak et al. 1986). Instead, the Maninka established Solo, as well as three other settlements to the east of Solo, near the Bafing River, outside the research area (Samak et al. 1986). This broader area, called the Bafing jamana (district), hosted at least 35 Maninka settlements by the late 1980s (Samak et al. 1986), when most of the Bafing was flooded following construction of the Manantali Dam (Figure 1, p. 52). The historical record is poor for the early period of Maninka settlement, until about 1890. Farmers established and abandoned several settlements in the research area other than Solo

26 (Figure 3, p. 54), primarily in order to improve access to farmland (Figure 5, p. 56). In many cases, however, settlements were established with a primary or secondary goal of improving their defensive situation during what was a time of political instability and warfare throughout Sudanian West Africa (Ajayi & Crowder 1985). Although jing (stone walls) were built to protect Solo and another large settlement occupied during the 1800s (Figure 2, p. 53), a more common practice of reducing the risk of violence was to establish new settlements in defensive locations (Mage 1868; Mollien 1820 [1967]; Noirot 1885; Park 1954 [1815]; Sanogo 1991; Tellier 1898), as elsewhere in nineteenth-century West Africa (Gleave 1966). In the research area, this practice meant that most settlements established prior to 1890 were built on or just below sandstone outcrops, which offered refuge. As a result, most settlements from this time had sandy soil rather than finer-textured soil (Figure 6, p. 57), which is most common farther away from sandstone outcrops (PIRT 1983). The research area was not a site of major warfare. Thus, unlike areas along the Bafing River to the northeast of Solo, which were essentially depopulated at the time of Eugne Mages 1868 visit, in the research area no Maninka settlements were abandoned directly as a result of warfare (Figure 4, p. 55). Oral history indicates that the possibility of war did lead to the abandonment of Solos second site after only c.50 years occupation, because its location was considered poorly defensible. There were probably other Maninka settlements in the research area before 1890 for which oral historical evidence does not survive, because hamlet settlement seems to have been practiced during this period. The best evidence for this comes from Mungo Parks journal from his second visit to what is now Mali, in 1805. Park probably visited Solos earliest location (Figure 2, p. 53), an unwalled village at the bottom of a rocky hill (1954 [1815]: 318), which

27 was abandoned soon afterward because of increasing concern about the hazard of rockfall from the adjacent sandstone outcrop. Less than about five miles west of Solo, Park (1954 [1815]: 317318) passed through the village of Gimbia, or Kimbia, then, after spending a night in Solo, observed several villages along the base of the cliffs immediately east of Solo. There is no oral historical evidence of these settlements. Gimbia was probably a hamlet occupied by people from Solo, located along the seasonal stream called Guimbaya k (creek at the place occupied by Guimba), where two hamlets named Guimbaya have been established and abandoned in living memory (Figure 3, p. 54). The villages to the east were also probably Solos hamlets, since the cliffs on the west side of the Bafing River along Parks route end about 5 km from Solos first location. Additionally, just west of the research area, Park (1954 [1815]: 296) passed through a very small village which he described as belong[ing] to another, larger village. This is how the village-hamlet relationship is described in the Maninka language, as explained below. While inconclusive, Parks journal suggests that hamlet settlement existed in the research area in 1805, and possibly also just to the west of this area, which is topographically similar and thus may have been equally protected from warfare. Parks journal does not suggest that hamlet settlement existed in the Bafing floodplain or other areas to the east, without sandstone outcrops. The expansion of European military authority throughout West Africa after about 1890 quelled warfare between African groups and initiated a period of peacealthough the colonial presence brought other risks, such as forced labor. As described above, shifting settlement probably existed before 1890 in the research area, especially near sandstone outcrops, but security and the conditions of colonial rule after 1890 (as described below) helped make shifting settlement an increasingly important means for farmers to respond to the new socieconomic

28 contexts of French colonialism and Malian independence. Current Maninka settlement practices are not timeless, but most strongly reflect twentieth-century socioeconomic and biophysical conditions. In order to understand how processes of socioeconomic and biophysical change affected settlement history after 1890, it is necessary to first describe Maninka settlement practice. Maninka settlement practice Settlement is a masculine practice for the Maninka. Men decide when and where to establish new settlements, as well as when to abandon settlements. Women, and to a lesser extent children, can exert informal influence on these decisions, but adult males have the only formal roles in settlement practice. The Maninka shifting settlement system allows farmers to exploit dispersed patches of arable soil in a socioeconomic environment where the costs of living distant from permanent villages are, for many people, low relative to the costs of remaining permanently in a village. The politics of hamlets. The primary reason for the establishment of 95% of occupied or abandoned settlements in the research area was the need for better access to farmland (Figure 5, p. 56). Koenig and Diarra (1998) also found this to be the case near Manantali, just north of the research area. Although a relatively large area of farmland surrounds Solo, it is limited. Solos traditional authoritiesthe dugutigi (village chief), dugukolo tigi (land chief), and other elder men acting as counselors (cf. Ciss 1970)decide how to allot Solos farmland to men and their families. Land rights in Maninka societies are primarily a function of age and historical primacy (Ciss 1970; Leynaud & Ciss 1978). In Solo, institutions of tenure strongly favor older men and men belonging to Solos founding family, whose oldest male is the chief. In principle, the chief holds rights to all land within Solos traditional territory, and grants usufruct for certain

29 areas to the male heads of Solos extended families. The head of each family manages most of the familys farmland as a foroba (large field), in which all family members work. The family head controls all produce from the foroba. Additionally, the family head may allot smaller fields to adult family members, both men and women, which they manage individually. These fields are the primary means through which women and young men living in Solo may earn money, because they retain all produce for their personal use. In all cases, however, the foroba takes precedence over smaller, individual fields (Leynaud & Ciss 1978). Thus, if a family head has use rights to less land or less productive land than he would like, he may decide not to allot individual fields to some or all household members. When such decisions are made, younger men and women are the first to lose access to farmland they can manage for their personal benefit. Maninka political and tenure institutions are essentially age-based hierarchies (Leynaud & Ciss 1978). The marginality of young men, regardless of their family connections, must be underscored in order to represent the importance of hamlet settlement in family development. Establishing or living in a hamlet is a normal aspect of household development in Solo, and virtually all people in Solo have lived in many hamlets during their lifetimes (Text Box 1, p. 50). Young men have relatively little influence on decisions about farmland allotment, and thus are often allotted less or less desirable farmland than they would like within a reasonable distance of Solo, usually about 3 km. As a result, their agricultural productivitythe source of their monetary income and their nuclear familys food securitydoes not meet its full potential, hindering household development in numerous ways. While land-hungry farmers do pursue the option of agricultural intensification to some extent through increased cultivation of secondary crops, the labor cost of substantial intensification far exceeds that of establishing a new

30 settlement at an unoccupied patch of farmland relatively distant from Solo (cf. Richards 1978; Stone 1996). Most families in hamlets are led by young, married men. Maninka shifting settlement centers on the social institution of the bugu (hamlet, farm), which allows young or politically marginalized men to increase their access to natural resources and space in a way that does not threaten the traditional political hierarchy. In a rural setting, the Maninka recognize two settlement types: bugu and dugu (village). Virtually all new settlements are bugu. Distinguishing villages and hamlets is often easy because most hamlets are small, geographically marginal, relatively undeveloped, and occupied by a few nuclear families headed by closely related men. However, some bugu are large, geographically central, well developed, and occupied by many nuclear familieslike most dugubecause the Maninka definition of these terms is political, not functional. Functionally, hamlets allow men to access more or better farmland than they can in their home village (Ciss 1970; Koenig & Diarra 1998). A man who wishes to establish a hamlet must negotiate with his familys head to decide how he will continue contributing to the extended family (usually by depositing a portion of his produce to the familys granary). As long as sufficient labor remains available for farming around the village, it is generally in the interest of a family head to allow younger members to establish hamlets because this increases the familys overall access to resourcesspecifically, distant patches of farmland. Hamlets allow Solos residents to efficiently exploit relatively small patches of farmland dispersed widely across the landscape. It is also in the interests of Solos traditional authorities to grant male residents permission to establish new hamlets because of their agroecological function. However, hamlets also pose the risk to Solos authorities that they may lose land rights around a hamlet. This can occur if a bugu grows large and endures over time, until its founding family, other residents, and

31 people in neighboring settlements consider it a dugu. Few hamlets ever attract more than a handful of nuclear families or last more than a couple decades, so this risk is small. Nonetheless, hamlets have a clearly secondary status politically: a bugu is considered to belong to a dugu. For instance, Sandiguila, a large settlement occupied for 80 years (until its eviction in 2006) was Solo ka bugu (Solos hamlet), and not a dugu. In daily life, this subordinate status means little to a hamlets residents because its men are free to manage the hamlet as they decide. However, all major decisions about land usethere is no criteria that makes a decision major, but an approximation is that any decision not anticipated within normal agricultural cycles is major belongs to Solos traditional authorities. For instance, in 1999 a man from a village south of Solo, in Kouroufing National Park, sought to establish a hamlet outside the park near Sandiguila. Solos traditional authorities permitted the man to build his hamlet, even though residents of Sandiguila thought he should not have been permitted to settle where he did. In general, political tensions between Solo and its hamlets are minimal, and men in both Solo and its hamlets regularly discuss land-use (and other) decisions as friends and relatives, in order to make the best possible decisions. Nonetheless, underlying these relationships is a clear political hierarchy that situates ultimate decision-making authority in Solo (cf. Ciss 1970; Leynaud & Ciss 1978). In the Bafing area, historical cases in which a village truly split, so that more than one resulting segment was recognized as a village, arose from intense disputes and social disintegration within the original village (Samak et al. 1986). Men choosing to establish new settlements accept as a cost decreased political authority and decreased access to the resources available in a village, but gain political autonomy and authority in their hamlets (although this is ultimately limited). Families or clans that have an unalterable, secondary political status in Solo have historically occupied long-lived, village-like hamlets because this situation offers them the

32 most authority and autonomy without causing social upheaval. For instance, the Dembl family, whose ancestor founded Solo, still dominates in the village, while the Kta family, whose ancestor arrived soon after Solos founder, occupied Sandiguila. The historical primacy of the Dembl family in Solo means that the Kta family will never be politically dominant, and thus will never have access to the best farmland around Solo. This situation is particularly galling to the current heads of Solos Kta family. When Sandiguila was evicted in 2006, its (male) residents decided to establish a new hamlet, rather than move to Solo, in order to maintain their autonomy. Overall, the costs of leaving Solo to live in a hamlet are minorespecially in a small, economically marginal village like Solorelative to the costs of agricultural intensification in existing fields, and the potential benefits of increased productivity in a new settlement. Site selection and management. While access to farmland is usually the primary criterion in selecting a settlement site, it is not the only one. Personal history, water availability, distance to other settlements and main paths, spirit activity, and other criteria are all considered in choosing a settlement site. Prior to 1890, the defensibility of a settlement site was another important criterion. Considering a possible settlement site in relation to the distribution of water sources (including groundwater), other settlements, paths, and various natural resources is a purely spatial analysis that is undertaken concurrently with the assessment of farmland in a site. Personal history and the activity of spirits are much more significant in constraining site selection. First, historical primacy is the main determinant of usufruct for the Maninka, and families retain settlement and cultivation usufruct for parts of the landscape in which their ancestors are the earliest remembered Maninka settlers (cf. Hopkins 1972; Leynaud & Ciss 1978; Samak et al. 1986). Physical features generally delimit such areas, which are often given

33 the name of the first settlement established in them. This name is also given to all subsequent settlements in a given area. For instance, Santankni is the toponym for a solon (large, basinshaped valley) northeast of Solo that has hosted at least six different settlements called Santankni, a name which was given to the first of these settlements by its founder c.1870 because it was the site of a santan tree (Daniellia oliveri) that was considered beautiful (k ni). The founders descendents retain unchallenged rights to renew settlement in this area even if no hamlets have been occupied in it for years, and in practice do not need to ask permission of Solos traditional authorities to do thisalthough in principle this is required. Conversely, individuals with family history in a particular area are constrained in their use of other parts of the landscape, because they must seek permission for settlement and cultivation in the other areas from both Solos traditional authorities and the usufruct-holding family. Often, the usufruct-holding family will grant permission to use only less-desirable patches of unused land, even if they have no plans for using more desirable patches, because they fear losing clear usufruct in the future. Men descended from Solos founder are its traditional authorities, and thus control the most and the best farmland near Solo. Second, the activities of jine (spirits) are extremely important in the success or failure of a settlement, as well as in farming, hunting, and all other activities (see Chapter 3). Many spirits, both good and bad, occupy the Maninka landscape. Several steps are taken to determine the suitability of a site for settlement with respect to spirit activities. To begin with, a man considering establishing a new settlement often seeks spiritual guidance by consulting a hunter who understands knyndiyon (literally sand children, referring to marks drawn in sand), an art of prognostication. Knyndiyon allows Maninka hunters, who are extremely knowledgeable about the spirit world (Cashion 1982; Ciss 1964), to foresee whether an action will benefit a

34 person, that persons health and material wealth, and the health and material wealth of that persons immediate and extended family. Based on the hunters findings, a man can decide if he should establish a new settlement, andafter having selected a potential sitewhether a site is suitable for settlement. In surveying possible settlement sites, those with obvious signs of malevolent spirit activitysuch as the abundance of huyuhiyo (Dombeya quinqueseta) or other plant species, or unusually warm or cool airare avoided. Sites exhibiting benevolent signs such as trees that have developed small root buttressesare favored. In sites without obvious, malevolent spirit activity, a man examines the soil nyama, which is the soils inherent healthfulness and is different from its fertility (see Chapter 3). By digging and refilling a small hole, and monitoring how this hole changes over several days (changes that are considered significant appear to relate to bulk density), a man, in consultation with others, can determine whether humans can safely occupy the site. These steps must be taken before establishing any settlement, even one that will be built within meters of an abandoned settlement that was a successful and healthful site for its occupants. Despite these precautions, a substantial number of settlements have been abandoned because bad spirits in a settlement site have caused many illnesses or deaths to residents (Figure 4, p. 55). Taking all these criteria into consideration, there are a limited number of areas across the landscape that are suitable for settlement, although most of these patches are unoccupied at any point in time. The limiting factor that reduces the number of possible settlement sites is soil patchiness (cf. PIRT 1983). Many parts of the landscape are not cultivable (see Chapter 3), and men can each identify several sites with high fertility, good soil nyama, and an absence of malevolent spirit activity that have never been farmed because the size of the arable patch is too small and too distant from other patches for cultivation to be practical. Thus, new settlements

35 tend to be located near abandoned settlements, and relatively distant from occupied settlements (Figure 7, p. 58). Once a site has been selected for a new settlement, its actual establishment entails much physical labor, which may be spread over several years. The first step of establishing a settlement is to clear an area for housing, mens work undertaken generally in the early dry season (January-March), after completing fieldwork and before seasonal surface-water sources have dried. Clearing vegetation in a settlement site differs from clearing farmland, because in settlements trees are cut in order to kill them, not simply to remove their crowns, which shade crops. Thus, trees are felled near the ground and near-surface roots are cuttwo acts which are completely avoided in clearing fields (cf. Nyerges 1989). Housing locations are often selected in relation to existing trees. Locations with very large trees are avoided because these represent hazards during windstorms, and may provide habitat for pest birds. Baobab (Adansonia digitata), shea (Vitellaria paradoxa), and locust bean (Parkia biglobosa) trees are spared, and each family spares about one medium-sized tree for shade, but all other woody plants are killed and removed (and not burned in place, as in fields). Depending on site-specific characteristics, the next step may be to dig a well, a task that men undertake toward the end of the dry season (May-June), when the water table is low. Most hamlets are located in sites with relatively high water tables, where it is easier to dig wells. However water may be availableon the surface or in a wellit is used to make adobe for building huts and granaries. Depending on resources available near a site, men may build with bricks or, more frequently, using bamboo wattles and daub. Women finish construction by sealing floors and walls with a hard-drying mixture of mud, the sap of various lianes, and cow manure, then painting designs on its outside using various mixtures of ash, soils, and ground siltstone. Construction techniques used in new settlements

36 often differ from those used in larger settlements, because techniques requiring a large number of workers cannot be used in situations where only one or two are present. Indeed, some settlements have been abandoned because the number of occupants became too low to sustain customary labor practices (Figure 4, p. 55). Once a minimal number of huts and granaries are builtin contrast to older settlements, men and women, and people and domestic animals may share huts for extended periods of time in new settlementswomen prepare the living area by clearing herbaceous vegetation and sweeping the soil, while men clear vegetation for fields. As time goes on, additional structures may be built, but, again, settlement establishment often occurs over several years, and young settlements are often undeveloped, minimalist places of shelter. Decisions to abandon settlements are usually made relatively easily. Most hamlets are essentially disposable because people plan to return to their residence in Solo, which they have maintained while living elsewhere. The mean length of occupation of settlements around Solo is 29.2 years, although the median length, 20 years, is lower because several settlements were or have been occupied for more than a century (Figure 8, p. 59). Hamlets are generally occupied for just one or two decades, but even villages and long-lived hamlets are regularly abandoned. In these cases, however, residents agree to move en masse to a new, nearby site that is, for some reason, more attractive than the older site. For instance: Solo has occupied three sites, separated by up to 3 km, since its founding >250 years ago; and from c.1926-2006 the hamlet of Sandiguila occupied five sites, separated by up to 2 km, near the site of the original Sandiguila, which was abandoned c.1894 (Figure 2, p. 53). Villages and long-lived hamlets are generally found in locations with abundant farmland and other resources, and can sustain long periods of occupation. As a result, these settlements become nodes in the local path network, and are crucial to maintaining communication between nearby settlements. Nonetheless, over time even

37 once-large settlements in resource-rich locations may dwindle and become liable to abandonment. In such cases, abandonment is not decided upon lightly, because the consequences may be significant for nearby settlements, especially in terms of decreased ability to communicate with more distant villages. While there is no central authority planning settlement location around Solo, the combined resource needs and use-rights of its occupants mean that settlements are widely dispersed across the landscape to reduce the rate of resource depletion in any given area, while certain significant locations with abundant resources are continuously occupied. Settlements are abandoned for many reasons, but limited access to farmland is the major reason why men decide to abandon settlements (Figure 4, p. 55). Hamlets are generally established on relatively small patches of farmland, access to which may decline as a result of population growth (due to the arrival of other families) or because there is only enough farmland to sustain one or two swidden cycles, insufficient time to restore fertility through fallowing. Whatever the reason, settlements sites that become unsuitable are often abandoned in stages over several years. First, women, children, and some men return to Soloor occasionally join an established hamletto establish new fields, while some men stay behind to exploit remaining fertility in existing fields. Subsequently, these men return to Solo but to begin with sometimes travel almost daily to the hamlet to manage fields, fallows, fruit trees, and multi-year crops (such as cassava, Manihot esculenta). Usually after about 4-5 years past occupants rarely visit abandoned hamlets, because new fields and gardens in Solo fully occupy themunless conditions in Solo have again proven unsuitable, in which case these people may have already established another new hamlet.

38 Over a lifetime, an individual will likely live in several settlements (Text Box 1, p. 50), generally returning to Solo for several years after abandoning one settlement and moving to the next, and returning to Solo permanently in old age. The labor and political costs of building and moving into several new settlements over several decades is less for most individuals than the costs of working marginal farmland for decades around Solo. Indeed, many of Solos current, traditional authorities lived in hamlets until their influence increased due to the death or illness of older, male family members (Text Box 1, p. 50). Other men, in less dominant families, experience similar increases in influence as they become elders, and, at some point, generally find that the benefits of this influence are greater than the costs of living away from Solo in a hamlet. Settlement history after 1890 The cost-benefit relationship of living in Solo versus a series of hamlets has been maintained through the 1900s in part by economic underdevelopment, which has maintained a low relative benefit of living in Solo. Combined with the patchiness of arable soil across the landscape and social differences that limit some mens access to farmland in specific parts of the landscape, Solos lack of development has reinforced the utility of shifting settlement as a means of exploiting the areas sparse socioeconomic and biophysical resources. Recently, however, political events and somewhat increased economic opportunities have increased the attractiveness of living in Solo. Adapting to colonial rule. During 1890-1960, farmers adapted to the socioeconomic context of colonial rule, even though the Bafing area was marginal to French Soudans economy and bureaucracy. Colonial rule affected settlement in two ways. First, the reduced threat of violence made farmers feel secure living distant from large settlements or settlements in

39 defensive locations (Samak et al. 1986). Similar changes in settlement practice have been widely reported in West Africa (Gleave 1966; Queant & de Rouville 1969; Sidikou 1974; Woodford 1974). Second, colonial rule brought no benefits to living in large settlements, but increased the actual or potential cost of doing so, through taxation and forced labor recruitment. In the colonial period, there were no public services in Solo such as a medical facility, water pump, cistern well, or school (although a primary school was founded c.1958)so living distant from Solo also carried minimal opportunity cost. People did not completely avoid paying taxes or supplying forced labor by living in small, remote hamlets instead of larger, more accessible villages, but in areas that were marginal in colonial geographies, people living in hamlets seem to have had the benefit of avoiding direct interaction with colonial authorities (cf. Hill 1953). In the case of Solo, direct interactions often brought costs that could not be passed on to or dispersed amongst other community members, such as taxation through confiscation of observed produce, or summons to provide forced labor given to observed individuals. Both of these factors provided incentive for a more dispersed settlement pattern than had existed before. Whatever the ultimate causes of increased hamlet settlement before 1960, the proximate cause remained, in all cases, improved access to farmland. Soil characteristics of settlement sites established during 1890-1960 suggest that the goal of agricultural production during this period was primarily grain production (Figure 6, p. 57). Most settlements were established in sites where silty to loamy soils predominated. The local grain staples, millet, sorghum, and fonio, thrive in these soils, which most other cropsfrom peanuts to yamstolerate even though other soil types suit them better. Export-oriented peanut production, which was important during the colonial period elsewhere in western Mali with better access to the Dakar-Bamako railroad (Hopkins 1972), did not concern farmers around Solo until the 1960s (cf. Kta 1972).

40 Settlements were abandoned for various reasons during 1890-1960. The most common reasons, however, were biophysical changes that made a site unsuitable for shelter, farming, or both (Figure 4, p. 55). The most important biophysical condition was soil fertility. Settlements founded to improve access to arable land lost all attractiveness if farmland fertility declined substantially. However, other biophysical changesfrom decreased water availability to increased pest and malevolent spirit activitymade some sites uninhabitable even if fertile farmland remained. Settlement following Malian independence. Following Malian independence in 1960, the Malian government sought to increase peanut exports by subsidizing production and processing (Kta 1972). This meant that peanuts became a crop with a guaranteed market and price, and most farmers in Solo focused their efforts on peanut farming. Most importantly, the government subsidized trucks to travel to all settlements, even hamlets, to purchase peanuts (Kta 1972). Thus, living in Solo lost the benefit it has otherwise always held as a village of having somewhat better access to agricultural markets via more frequent visits from traveling merchants, and a greater number of young, male residents who regularly carry produce to commercial markets, dozens of kilometers away. As a result, beginning in the early 1960s, many new settlements were established in areas with sandy soil, in which peanuts grow very well (Figure 6, p. 57). Most of these sandy-soil settlements were abandoned in the early 1980s, when state subsidization of peanut farming ended (Figure 3, p. 54). Since then, settlements have been established in sites with a wide range of soil environments, including many with silty soil, because farmers have renewed their focus on subsistence and grain production. For the period since 1960, settlements have been abandoned for several reasons (Figure 4, p. 55). Biophysical causes, especially soil fertility decline, remain very important. Settlement

41 abandonment due to water shortageeither because the water table has descended, or surface water sources have driedhas become more common since 1960, and may be linked to the longterm drought observed across Sudanian West Africa (cf. Nicholson 1996). Social causes have become increasingly important in settlement abandonment (Figure 4, p. 55). Throughout the 1900s some settlements were abandoned because the departure of some residents, or the abandonment of a nearby settlement, meant that remaining residents felt socially isolated. Such loss of community has been the most important social cause of settlement abandonment since 1960 because in the late 1980s over thirty settlements in the impondment zone for the Bafing Reservoir were relocated north of Manantali Dam (Figure 1, p. 52). Although little of Solos traditional area was flooded, most settlements in the Bafing area disappeared, representing a loss of community for Solo as a whole. As a result, most of Solos occupants, including nearly all of its hamlets, decided to abandon their settlements and move to the resettlement zone. Solo and several of its hamlets were never completely abandoned, though, and since the early 1990s many people have left the resettlement zone to return to the Solo area, and live closer to family and friends. Another increasingly important cause of settlement abandonment has been eviction by political authorities. The Malian nature conservation directorate has carried out all evictions since 2004, with the intention of reducing wildlife habitat loss in Kouroufing National Park. Since only part of the research area is in the national park, these evictions have been accompanied by several cases of settlement establishment outside park boundaries. Since the mid-1990s, several new hamlets were built on the plateau north of Solo and the park boundary out of fear of eviction, even though BBR authorities had not at that time publicly presented their ideas on eviction. In 2004-2005, Solos three hamlets in the national park were finally evicted,

42 and most of the affected people built a new hamlet on the plateau, near a small, preexisting hamlet. All of these hamlets are in an area near the edge of the plateau, near where previous settlements had been established in easily defensible sites. Historically, most families who abandoned hamlets did not move directly to another one, but returned to Solo for some several years or more before moving again to a hamlet. These returns to Solo renew social contacts and provide new opportunities unavailable in hamlets. Solo is not disintegrating, but during the early stages of household development most men have substantial incentive to leave Solo and take advantage of hamlet settlement. In recent years, living in Solo has offered somewhat more benefit than in the past, which may in the long term reduce incentives for hamlet settlement. First, there is considerable uncertainty about the new hamlets north of the national park, since this area is part of the proposed buffer zone. Solo is legally a village, and thus appears to face little threat of eviction. This fact attracted several evictees from hamlets that had been in the national park, even though these people could have probably gained access to more or better farmland by joining the new hamlet on the plateau north of Solo. Second, the few socioeconomic resources available in Solo are becoming increasingly attractive because the Bafing area has gained increased attention from regional development and conservation projects. While there has been little real effect of these efforts in Solo, small developmentsgovernment appointment of a teacher for the community-funded school, increased opportunities for employment in conservation and research projects, the establishment of a grain bankhave caused a small number of men to choose to remain in Solo rather than move to or remain in a hamlet (Figure 4, p. 55). The increased importance of distance to main paths as a cause for settlement abandonment also indicates the increasing importance to Solos farmers of access to larger-scale socioeconomic networksprovided, for instance, by visits from

43 traveling merchants, medical practitioners who vaccinate children, and political campaigners, all of whom travel only on main paths. Solo, although economically and geographically marginal in southwestern Mali, has better access to these large-scale resources than its even more marginal hamlets, and thus has attracted many men to reside again in Solo, rather than a hamlet. Nonetheless, settlement continues to be a fluid process, and observation of settlement pattern at any point in time remains only a snapshot that becomes inaccurate with time (Figure 3, p. 54). For instance, of the five Solo hameaux PREMA identified in 1996, one was abandoned within two years and three were evicted in 2004-2006, and one new settlement has been established. Furthermore, three hamlets had been abandoned in the five years prior to 1996. It is inaccurate to interpret or represent Maninka settlements as permanent, fixed points in a distribution to which new points are continuously added over time. Settlement and conservation in the BBR Bugu, hameau, hamlet; dugu, village, village: These trios have essentially the same physical meaning. Indeed, the pattern of Maninka settlement is not difficult to discern: dispersed, small, young settlements outnumber larger, older ones. However, the functional meaning of these terms differs. The process of settlement expansion implied by the French terms hameau and village in conservation policies differs from the process of settlement shifting experienced by Maninka farmers. Misinterpretation of settlement pattern in the BBR has created a political environment in which economically and politically marginal men and their families marginal locally, regionally, and nationallycan be effectively punished for a humanenvironment interaction that does not exist, or at least not in the manner in which it is represented.

44 Habitat loss to farming, livestock herding, logging, road building, and mining is probably the greatest medium- to long-term threat to biodiversity resources in the BBR, but the actual processes of habitat loss are poorly understood (Duvall et al. 2003). While settlement is inextricably linked to these other land uses, settlement is a distinct land use that is not a direct cause of habitat loss in the BBR. Current conservation policies that have evicted and prohibit hamlets are blunt and inefficient means of reducing habitat loss that will likely fail in the long term for two reasons. First, these policies seek only to change settlement patterns, and neglect the reasons why men establish new settlements. The policy of restricting village populations to territoires des villages (Caspary et al. 1998) will only intensify the socioeconomic and spatial processes that make living in hamlets more attractive to many men than living in villages. The policy of prohibiting the establishment of new settlements will increase hardships for economically and politically marginal men and their familiesassuming, unreasonably, that this policy can be enforced and that these men will not simply establish a new settlement elsewhere. A much more effective way to decrease the number of hamlets in the BBR and increase the proportion of men who choose to live in villages would be to increase the range, availability, and quality of socioeconomic resources in villagesincluding schools, roads, water pumps, and medical facilities. On a broader scale, economic policies that decrease the costs of agricultural intensificationsuch as subsidizing the cost of chemical fertilizerwill also decrease the rate at which hamlets are established (Koenig & Diarra 1998). By intensifying competition for farmland and increasing economic hardships for many people, current, coercive conservation policies will diminish local support for conservation goals and activities, and preclude conservation success in the long term.

45 The second reason that current policy toward settlements will fail to meet conservation goals is that it neglects the geography of biodiversity and settlement in the BBR, and will intensify human environmental impacts in parts of the landscape where biodiversity is highest. The boundaries of the BBRs national parks were drawn with limited knowledge of the distribution of biodiversity resources. Although these national parks protect areas with relatively low human population density and the highest local densities of several antelope species (Caspary et al. 1998; Duvall & Niagat 1997), they protect negligible amounts of the highly biodiverse habitats characteristic of sandstone outcrops, in which chimpanzees are most frequently observed (see Chapter 6). These habitats receive little protection via conservation law enforcement, but they receive passive protection because they occur in nearly inaccessible locations that people rarely visit (Duvall 2001). However, in areas with relatively high human population density these habitats are used more frequently for logging and hunting, which has contributed to chimpanzee range contraction during the last twenty years (Duvall 2000; Duvall et al. 2003). Most people evicted from hamlets in the northern part of Kouroufing National Park have established new settlements near the edge of the sandstone outcrop just north of the national park. Assuming that the prohibition of new settlements in the BBR will continue, more men from nearby villages will decide to establish or join hamlets along this outcrop. The increased population of this area will increase the environmental impacts of humans on outcrop habitats, and the vacated parts of the BBR are not equally valuable for biodiversity, or chimpanzee, conservation. Evicting hamlets may guarantee a low population density in the BBRs national parks, but in the long term this may increase threats to the reserves nationally and internationally most important biodiversity resources. Furthermore, decreasing or eliminating

46 possibilities for hamlet settlement will likely encourage men to farm increasingly marginal farmland around existing settlements on an increasingly permanent basis, and thus lead to permanent habitat conversion (cf. Koenig & Diarra 1998; Nyerges 1989). While conservationists have accused Maninka farmers of using land without any planning beforehand (Caspary et al. 1998: 77), current policies on settlement in the BBR seem fairly nearsighted. More effective conservation policies for the BBR would take advantage of Maninka settlement practice. Indigenous Maninka residents will need to modify their resource use, especially hunting, if biodiversity conservation is to succeed, but the dispersed, shifting pattern of Maninka settlement should be seen as a conservation resource. Dispersed settlements mean more dispersed human environmental impact, and also spatially more uniform and strategic surveillance for illegal poaching, logging, and other activities, especially since most large settlements are located on main paths. Conservationists have recognized the potential of local residents to contribute to conservation goals in this way by establishing surveillance committees in many villages in the BBR (cf. Caspary et al. 1998). However, such surveillancewhich happens regardless of conservation efforts because people monitor the areas around their villageswill contribute to conservation only if people in hamlets are empowered to have an interest in reporting what they see to law enforcement officials. Settlement policies, enforced by these same officials, that result in hardship remove this interest. Conclusion Human-environment geography must recognize rural settlement as a distinct land use. Human-environment geographers, using the analytical tools of cultural and political ecology, have contributed significant theoretical and practical knowledge to resource management in rural, agrarian landscapes through careful examinations of other rural land usesespecially

47 agriculture, pastoralism, and conservation. However, the failure to study settlement as a distinct land use has limited human-environment geography in two key ways. First, settlement processes occur at specific spatial and temporal scales (Stone 1996), and the failure to recognize settlement as a distinct land use has meant that scales of observation used in human-environment geography have often been inadequate to observe significant aspects of resource use in rural, agrarian areas. Many human-environment geographers have approached questions of resource use from the perspectives of cultural ecology and political ecology. Many cultural ecologies descend methodologically and theoretically from important works such as Conklin (1961) and Boserup (1965). These works, and many of their descendants, focus on agricultural practices at the scale of individual plots or settlements, and thus fail to recognize how agriculture relates to settlement processes that occur over landscapes (areas of tens to hundreds of square kilometers). On the other hand, political ecologies of resource use, which explicitly recognize the significance of processes operating at scales broader than communities (Robbins 2004; Zimmerer & Bassett 2003), have generally jumped over the landscape scale to focus on national, regional, or international processes. Human-environment geographers have focused on landscape scales mainly just in the context of pastoralism, because the movement of people and livestock across landscapes is obviously crucial to this land use and abundantly obvious over even brief periods of observation. Mobility is also inherently important in settlement in rural, agrarian landscapes, but this mobility is often not apparent except over decades. Settlement pattern spatially structures resource use across landscapes (Chisholm 1979; Stone 1996), and accurate understanding of resource use requires understanding of settlement processes that unfold over decades and landscapes.

48 Second, by focusing on land uses other than settlement, human-environment geographers risk simplifying or overlooking settlement processes, and giving the impression that spatial patterns of settlement accurately substitute for knowledge of these processes. Too frequently, analysis of rural settlement in human-environment geographies is limited to statements about the spatial pattern of settlement, reflecting a long tradition in cultural geography that relates the form and distribution of settlement to various social, economic, cultural, and physical geographic factors (cf. Hill 2003). However, different biophysical and socioeconomic environments can create different settlement processes that lead to similar settlement patterns (Stone 1996). For instance, Ruthenberg (1980: 31) suggests that shifting settlement may develop where shifting cultivation is practiced and, over time, [t]he cultivated plots move slowly away from the previous clearing and the vicinity of the hut. [Thus,] the cost of transportation increases[]. Beyond a certain distance, it becomes advantageous to build a new hut near the field instead of carrying the harvest such a long way. This process of shifting settlement is certainly accurate for many places and times, possibly including many areas where there is frontier-style settlement expansion (Netting 1993: 223). However, transportation cost does not lead to shifting settlement in Malis Bafing area. Instead, the patchiness of arable land, and differences between individuals in their abilities to access good farmland near villages, creates conditions in which shifting settlement proves beneficial to most families when considered over a decadal timescale. Failure to recognize settlement as a distinct land use whose observed patterns have distinct and variable formative processes has limited the ability of human geographers to accurately and precisely understand land use in agrarian, rural landscapes.

49 Text box and figures for Chapter One

50 Text Box 1. Hamlets and young families. Hamlet settlement is an integral part of household development for people in Solo, as illustrated by the personal histories of Mbakuru, a 19-year old woman, and her father Jigiba, a 48-year old man. The names of these people have been changed to protect their privacy. In late 1987, Mbakuru, her parents eldest, was the first child born in New Solo, where many of Solos residents resettled after construction of the Manantali Dam. Prior to this, Jigiba and his wife had been living in a hamlet south of Solo, which he had established in 1981, near an abandoned hamlet, because he couldnt get good fields in Solo. In 1986 Jigiba decided, along with most of Solos men, to move their families to New Solo, in the resettlement zone north of the dam (Figure 1, p. 52), in order to remain near other resettled villages. Jigiba soon became frustrated by poor farming conditions at New Solo, and in 1990 moved his family back to Solo, where his father had become dugutigi (chief). In 1992, Jigiba and two cousins decided to establish a new hamlet south of Solo, where they thought they would have greater agricultural productivity, based on Jigibas having lived in two nearby hamlets (that his father had joined in the 1950s and 1960s) as a child and young man. Farming was successful in the new hamlet, but it was abandoned in 1997 primarily because it was too far from a main footpath: there were few visitors, and it was too difficult to transport produce to Solo. In 1999, Jigiba began planning a new hamlet, but in 2000 his father died, giving him increased responsibility and authority within his extended family. One of his cousins with whom he had established the hamlet in 1992 became dugutigi. Additionally, Jigiba has found intermittent employment in Solo working as a guide for researchers and visiting Peace Corps Volunteers, and does not wish to lose this opportunity by moving to another hamlet. He has probably made the transition to fixed settlement in Solo.

51 After living in several hamlets with her parents and younger siblings, Mbakuru was married in June 2004, and moved to Kama, her husbands village, about 20 km west of Solo. In 2005, however, her husband decided to move with her to a hamlet founded by his cousin in 2002, where they now live (except during the dry season, when they return to Kama). Mbakurus first child, a daughter, was born in the hamlet in late 2005.

52 Figure 1. Western Mali, showing location of Solo and the Bafing Biosphere Reserve. Abbreviations: BZ=buffer zone for Bafing Biosphere Reserve (BBR); KNP=Kouroufing National Park, part of the BBR; WNP=Wongo National Park, part of the BBR.

53 Figure 2. Map of the research area. Only settlements named in the text have been shown. The abandoned settlement sites shown for Solo, Sandiguila, and Santankni are numbered in the order of their establishment.

Santankni sites 1 2 4 3 5 6 Solo 1 Solo 3 Solo 2 1 6 2 4 3 5 Guimbaya Sandiguila sites sites

1 2

Primary footpaths Seasonal streams Bafing Reservoir Kouroufing National Park Clif Occupied settlements Settlements evicted 2004-06 Settlement established 2004 Abandoned settlements mentioned in text Walled settlements, 1800s Study area

5 km

54 Figure 3. Distribution of settlement sites in the research area. Only settlements for which there is oral historical evidence are shown. Four time periods are represented: 1) pre-Maninka is the period up to and including the Maninka occupation of the area, which occurred >250 years ago; 2) early Maninka is the period of Maninka occupation of the research area up to 1890; 3) 18901960; and 4) 1960-2006. Black points show settlements established during each time period; open squares show settlements abandoned during each time period.

Latitudinal distance (meters)


early Maninka



Longitudinal distance (meters)

55 Figure 4. Causes of settlement abandonment. Wide, gray bars show primary causes; narrow, black bars show secondary causes. Total number of settlements abandoned (n) indicated per time period. Data from oral historical interviews. Time periods as described in Figure 3 (p. 54).

56 Figure 5. Causes of settlement establishment. Wide, gray bars show primary causes; narrow, black bars show secondary causes. Total number of settlements established (n) indicated per time period. Data from oral historical interviews. Time periods as described in Figure 3 (p. 54).

Causes for establishment Improve farmland access Outside national park Near water source Reoccupy past residence Good spirits in site Defensive location Unknown cause

Number of settlements 12 0 4 8 12 33 39

4 15

1960-2006 (n=35)

1890-1960 (n=39)

early Maninka (n=16)

preManinka (n=2)

57 Figure 6. Soil texture at settlement sites. Time periods as described in Figure 3 (p. 54).

58 Figure 7. Spatial pattern of settlement sites. Values for Ripleys L >0 indicate attraction, while values <0 indicate repulsion. Thus, settlement sites display attraction at distances <2.3 km and repulsion at distance >2.3 km. Increasingly positive/negative numbers indicate stronger attraction/repulsion. Observed attraction indicates sequent occupation of preferred habitat patches by multiple settlements over time, while observed repulsion indicates: a) the patchiness of preferred habitat, and b) the likelihood that nearby, contemporaneous settlements are separated by a minimum distance, since the field-to-settlement distance many men consider reasonable is c.3km.

59 Figure 8. Histogram of temporal lengths of occupation for settlement sites.

60 Chapter Three: Folk taxonomy of physical geographic terms used by Maninka farmers in southwestern Mali

Abstract This paper analyzes the nomenclature and taxonomy of physical geographic terms in the Maninka language as spoken in the Bafing area of southwestern Mali. Its purpose is to understand the content and structure of this particular body of local knowledge, and to compare Maninka physical geographic knowledge to that of other cultural groups. The research is based on participant observation and ethnographic interviews. Main findings include: 1) The Maninka conceptually bifurcate the biophysical environment based on whether natural resources contained in observed physical features are openly accessible to all humans. Features that are owned or otherwise possessed by humans or spirits are not accessible, and are classified according to physical criteria and the abstract quality of possession. Openly accessible features are classified based on physical criteria. 2) The main criteria used for classifying openly accessible features are: hydrology, topography, ground characteristics, vegetation, and microclimate. Physical features are classified by these criteria alone, or by a sixth criterion representing a synthetic view of all resources present at a given site. 3) Many classificatory criteria reflect evaluation of resources based on use-value in the context of Maninka agricultural practice. 4) Maninka natural resource classification is similar to that reported for related and other cultural groups. However, culturally specific classifications of locally diverse or highly valued resources are embedded within the cross-culturally similar, broad framework. This paper concludes that greater attention should be given to the broad conceptual context of physical geographic terms or concepts reported in ethnoscientific analyses of local knowledge systems.


Keywords indigenous knowledge; ethnoscience; folk taxonomy; local knowledge; ethnopedology

Introduction Indigenous farmers and pastoralists successfully manage diverse ecosystems using sophisticated knowledge of the biophysical environment. Unlike technical scientific knowledge, local knowledge1 is mediated through everyday language; terms that carry ecologically significant meanings often carry meanings in other, seemingly unrelated, contexts. Layered meanings, often overlooked by outsiders, create the context in which local knowledge gains and retains meaning for its users (Agrawal 2002). Yet outsiders often emphasize limited aspects of local knowledge systems in order to underscore practical applications these may have in natural resource management. The practical value of local knowledge is not in question, but overemphasizing its potential applicability in limited contexts leads to incomplete characterization of local knowledge of the biophysical environment2 (Agrawal 2002; Scott 1998).

Following WinklerPrins (1999), I prefer local knowledge to indigenous, traditional, or folk knowledge because a key attribute of these types of knowledge is their geographically limited extent. Local knowledge also implies nothing about the history of knowledge or of those who retain it. I use folk taxonomy because it is an established technical term without synonyms. I consider terms of the form ethnoscience to denote studies of local knowledge rather than the local knowledge itself. 2 Biophysical environment refers to all biological and non-biological, physical features in an area. Natural environment or environment sometimes carry this meaning, but the former excludes anthropogenic features, and the latter lacks specificity. Physical environment refers only to non-biological features. Biophysical environment differs from biospiritual environment, a portion of Maninka geographical knowledge, discussed below.

62 During the last fifty years, many researchers have employed an ethnoscientific approach to studying local knowledge of the biophysical environment, which entails analysis of particular aspects of local knowledge systems comparable in referential extent to specified technical scientific fields. Ethnobotany, ethnozoology, and ethnopedology have received the most attention (e.g. Balick & Cox 1996; Barrera-Bassols & Zinck 2000; Berlin 1992; Cunningham 2001; Medin & Atran 1999; Winklerprins & Sandor 2003). A few have described local knowledge of climate (Goloubinoff et al. 1997; Osunade 1994; Ovuka & Lindqvist 2000), while others have studied local knowledge of habitat variation (Fleck & Harder 2000; Frechione et al. 1989; Osunade 1988; Osunade 1987; Shepard et al. 2001). Very few researchers have analyzed how knowledge of the biophysical environment as a whole is structured in local knowledge systems (Barrera-Bassols & Zinck 2003b; Goodenough 1966). This is the referential frame Blaut (1979) suggested for ethnogeography, the study of how cultural groups perceive variation in the biophysical environment. As an ethnoscience, ethnogeography is underdeveloped3, even though Blauts holistic approach avoids a priori compartmentalization of indigenous knowledge into categories with limits determined largely by imposed correspondence with technical scientific fields of study. Blauts concept of ethnogeography is similar to ethnoecology, which Barrera-Bassols and Toledo (2005: 11) define as the study of how nature is perceived by humans through a screen of beliefs and knowledge, and how humans, through their symbolic meanings and representations, use and/or manage landscapes and natural resources. Ethnoecology emphasizes the complex layering of local environmental knowledge, from spiritual belief systems that guide

Unfortunately, most subsequent uses of ethnogeography do not follow Blaut (1979), but refer to descriptions of the distribution of cultural groups.

63 resource use, through bodies of knowledge underpinning resource use, to practices of resource use (Barrera-Bassols & Toledo 2005; Barrera-Bassols & Zinck 2000). The desire to maintain local knowledge in context, and ultimately in situ, strongly and explicitly motivates ethnoecological research. The ethnoecological approach holds great promise for advancing understanding of local environmental knowledge systems. However, ethnoecology has continued, albeit subtly, to compartmentalize local knowledge according to technical scientific criteria. The desire to identify practical applications of local knowledge has also been a strong and explicit motivation for ethnoecological research (cf. Barrera-Bassols & Toledo 2005). This goal is arguably at cross purposes with the goal of maintaining local knowledge in sociocultural context (Agrawal 2002; Scott 1998), and has led ethnoecologists to privilege certain research questions over others. Specifically, a central theme in ethnoecology has been the comparison of local knowledge to specific domains of technical, scientific knowledge (Barrera-Bassols & Toledo 2005), especially soils (cf. Barrera-Bassols & Zinck 2000; Winklerprins & Sandor 2003). Yet ethnoecologists have not shown and do not argue that technical, scientific knowledge provides a panenvironmental standard by which other knowledge systems should be assessed. Comparison of local knowledge systems across cultures and environments has not been a central theme in ethnoecology (Barrera-Bassols & Toledo 2005). Do humans similarly perceive variation in the biophysical environment across environments and cultures? Many ethnolinguistic groups have highly detailed knowledge of specific environments, and may have highly detailed classifications of features in those environments. Folk taxonomies of plants and animals show, in many cases, universal similarities (Berlin 1992), although biophysical variation between environments and socioeconomic

64 variation between ethnolinguistic groups lead to differences in the detail with which people distinguish biophysical features (e.g. Birmingham 2003; Voeks 1998). Does local knowledge of other biophysical featuressuch as landforms, water bodies, or precipitationsimilarly correlate to environmental and cultural variation? Geographers are well equipped to address these questions, which have been previously considered mainly by anthropologists. Cultural anthropologists have explored how humans perceive organisms across cultures and environments, and have found pan-environmental similarities in folk taxonomies of plants and animals (Berlin 1992; Brown 1984; Holman 2005). Geographers and others have used the methods of folk taxonomy to study local soil knowledge (e.g. Williams & Ortiz-Solorio 1981), but other physical features have not been subject to such analysis. Studies of local soil knowledge are considered in more detail in the discussion section, below. Evolutionary anthropologists have proposed that humans prefer habitats that resemble African savannas because this is where humans evolved (Orians & Heerwagen 1992). If this is the case, comparing local systems of physical geographical knowledge should indicate pan-environmental similarities in concepts of vegetation structure, if not other biophysical features. Analysis of local environmental knowledge using folk taxonomy is important for identifying the conceptual bases of geographyspecifically, how culture may or may not constrain individual perceptions of the biophysical environment (Blaut 1979). The present paper contributes to our understanding of cross-cultural and -environmental variation in physical geographic knowledge by analyzing the content and conceptual structure of Maninka physical geography in southwestern Mali, and comparing this with published information on other local knowledge systems. The Maninka belong to the wider Manding culture, which consists of farming societies living throughout West Africa, mainly west of Ghana

65 and Burkina Faso (Figure 1, p. 92). Analysis of the nomenclature and taxonomy of Maninka physical geographic terms enables comparison of Maninka physical geographic concepts with those of other Manding groups. Cultural ecological research on several Manding groups is substantial, including recent works on Maninka (Laris 2002), Kuranko (Fairhead & Leach 1996; Nyerges 1989; Richards 1985; Richards 1995), Mandinka (Carney 1996; Carney 1991; Schroeder 1999), and Jula (Bassett & Koli Bi 2000). There are also many older works on these and other groups, particularly Bamanan farmers in Mali. Nonetheless, it is unclear how widely these groups share physical geographic knowledge described for specific groups in limited areas. Manding groups have strong historic, linguistic, and cultural links (Hodge 1971), but occupy a wide range of biophysical environments, from the Guinean rainforest edge to the Sahel, and from mangrove-dominated seashores to woodland-dominated uplands (Figure 1, p. 92). The goals of the present paper are to: a) inventory Maninka knowledge of physical geographic features, including processes that link features; and b) determine how this knowledge compares with that of other Manding groups. Understanding how Maninka physical geographic knowledge compares with that of culturally related groups in different environments provides a basis for identifying and understanding pan-environmental conceptions of variation in the biophysical environment. Research setting Field research was conducted January-December 2004, in Solo village in the Bafing area of southwestern Mali (Figure 1, p. 92). Solo was established 400-500 years ago (Samak et al. 1986). About 250 people live in Solo, subsistence farmers who rely on rainfall to grow millet, peanuts, and several minor crops. Residents also hunt, fish, collect honey, and gather wild plants to supplement their diet or supply local markets (cf. Horowitz et al. 1990; Samak et al. 1987).

66 Farmers follow a complex land management system to maximize crop security under constraint of the regions variable precipitation regime (Koenig et al. 1998; Laris 2002; Samak et al. 1987). Most residents have lived in Solo their entire lives, although many spend rainy seasons in small farming hamlets dispersed in an area of 183 km2 around Solo, where residents have traditional usufruct recognized by neighboring villages. This customary tenure is not recognized in state laws. In practice, Solos traditional authorities, principally the dugukolotigi (land chief), make decisions about allotting land for new fields or hamlets (Ciss 1970; Samak et al. 1987). Individuals make decisions about the use of particular resourcessuch as trees for fuelwood, or wildlife for meatwithout needing the approval of traditional authorities, although the dugutigi (village chief) often must resolve disputes between individuals about use rights. The Bafing landscape is topographically complex. Sandstone plateaus rise 100-300 meters above undulating plains and valleys that culminate in deep gorges incised into the plateaus (Michel 1973). Scree slopes form below cliffs along the plateaus. Several sandstone formations occur in the area, each having distinct color, hardness, and other characteristics; various fine-grained sedimentary rocks interpose the sandstone layers (Varlet et al. 1977). Sandy soils dominate (Dames & Moore 1992). Ferricrete crusts occur widely in the landscape. The edges of these crusts are steep and gravelly, while their upper surfaces may be barren, exposed hardpans or shallowly covered by silty to clayey soil (Michel 1973). Dolomite intrusions outcrop to form steep, rounded inselbergs surrounded by silty soil. Vegetation variation corresponds to hydrological and edaphic variation (Breman & Kessler 1995; Lawesson 1995). The area is in the broad band of Sudanian woodland that extends across West Africa (White 1965; White 1983). Different woodland associations occur in sandy and silty soils (Nasi & Sabatier 1988). Grassland or wooded grassland occurs in sites with

67 shallow soil. Sites with deep soil are favored for agriculture; nearly all have been farmed in the past two centuries (see Chapter 4). In sites that have been undisturbed for several decades, forest vegetation forms, dominated by woodland species. Along permanent or seasonal drainages, gallery forest occurs, dominated by species characteristic of more humid climate zones (Duvall 2001; Lawesson 1995). Hydrological resources are not diverse (Dames & Moore 1992). The most important source of water for humans is hand-dug wells. Rainfall is highly seasonal, with nearly all precipitation falling between July and October (Barth 1986). There are two semi-permanent creeks, many seasonal drainage channels, and several permanent springs in the 183 km2 research area. The original course of the Bafing River, the main tributary of the Senegal River, is about 20 km from Solo, outside its traditional territory. Dammed in 1989, the Bafing River now forms a large reservoir whose shore is about 8 km from Solo, on the edge of its territory. The reservoir is becoming an important transportation route and valuable fishing site, but for most residents it remains beyond their normal sphere of experience. Research methodology Orthography and language. This paper uses terms in English, French, and Manding. Maninka (or Malink), Bamanan (or Bambara), and Mandinka (or Mandingo) are closely related dialects of the Manding (or Mand) language, spoken widely in West Africa (Bird 1982). Differences between dialects are mainly systematic changes in pronunciation, and reflect historic or geographic separation of Manding populations (Bird 1982; Derive 1990). Maninka words collected during field research are written in bold italics; spelling follows Birds (1982) linguistic analysis and orthographic conventions. For clarity, only singular Maninka nouns are provided, even if a plural is given as a gloss or implied by the English

68 context. The plural marker for most Maninka nouns is lu (Bird 1982). Covert concepts are given English names and written in brackets, such as [slope]. Previously published terms are italicized in quotes, and written as in the cited sources. The meanings of English technical terms are from Thomas and Goudie (2000). Glosses for Maninka words come from field research. The meanings of previously published words come from the cited sources. Data collection and analysis. Participant observation offered opportunities to learn the use of Maninka terms, while ethnographic interviews clarified observations. I lived in Solo January-December 2004, and participated in about 600 hours of relevant conversation, including interviews. Conversations, all in Maninka, were conducted while hiking with hunters or working with farmers. Interview data came from 35 informants, male and female, aged 13-80. I began research by seeking names for physical geographic features, particularly landforms, soils, and atmospheric phenomena. Once I had developed a functional vocabulary of physical features, I shifted my focus to understanding relationships between features, and the taxonomy created by these relationships. In interviews, I asked names for physical features my interviewee and I observed together, then asked how observed features differ from similar ones I knew from previous experience. I also asked informants to indicate physical features whose names I knew, but whose forms I did not. When informants showed me features, I sought to determine precisely what they were indicating, and to collect names for related features. I used specific question formats, especially triadic and dyadic comparisons (Cotton 1996), to assess conceptual relatedness. Data analysis was qualitative. First, listening to conversations revealed grammatical and semantic classes. Basic grammatical classeslike noun, verb, locative noun, possessive, and

69 descriptive phraseoften suggested broad conceptual categories, while semantic analysis revealed polysemic terms and covert taxa (Berlin et al. 1968; Berlin et al. 1973; Conklin 1962; Kay 1971). Second, informants generally responded to comparative questionssuch as is X different from Y?with phrases whose meanings in terms of similarity ranking proved comparable between individuals. To express high to low similarity, informants said: wolu kr be kilin (their meaning is one), wo be kilin (it is one), wolu ka muno (they are similar), wo te kilin (it is not one), and wolu te muno (they are not similar). Repeated instances of informants using a single phrase in response to specific comparisons clearly indicated conceptual relatedness. Finally, after about 400 hours of conversation and interviews, a taxonomic structure was developed in multiple iterations to represent the relatedness of Maninka concepts (cf. Berlin 1992; Brown 1984; Kay 1971). Discussions with key informants tested whether proposed taxonomic relationships accurately reflected their knowledge of these concepts. Based on these discussions, proposed taxonomic structures were changed or retained. Technical scientific equivalents of Maninka soil and vegetation categories come from a concurrent study of vegetation characteristics (see Chapter 5). Soil texture was identified using manual texturing (Midwest Geosciences Group 2003) of samples from 217 sites where informants provided a Maninka name for the sampled soil unit. Woody vegetation was sampled using 0.1-ha plots (Phillips & Miller 2002) at 206 sites where informants provided a Maninka name for either vegetation or land cover. According to tree stem density and crown height (Lawesson 1995: 24), vegetation was labeled forest, woodland, or wooded grassland. Rock types were identified from descriptions in Varlet et al. (1977) and Groupement Manantali (1979). Results

70 Broad concepts. The Maninka biophysical environment is conceptually bifurcated into [the biospiritual environment] and [the physical environment] (Figure 2, p. 93). This paper focuses on [the physical environment]. [The biospiritual environment] comprises all beings biological or spiritual things that die and are susceptible to illnessand their possessions. The four categories of beinghadamadn4 (humans), jine5 (spirits), [animals], and [plants]are not clearly separable for several reasons. First, most informants consider humans a type of animalspecifically, a type of subo (mammal). Many animals, especially large vertebrates, share with humans the characteristic of having a ja (soul), spiritual power that remains after an individuals death. Hunters must propitiate the souls of animals they kill to avoid retribution (Cashion 1982; Ciss 1964), but the need to respect an animals ja diminishes as body size decreases. Second, subaga (sorcerers) can change forms freely between human and animal. Some informants also believe sorcerers can also transform between human and plant forms. Third, jine (spirits), which are generally dangerous, can change forms to look like humans, animals, or plants. Thus, humans must cautiously interact with other beings because these may not be what they seem. Most human activities represent acts meant to protect against, or gain the favor of, spirits (cf. Brun 1907; Lem 1948). Jine also interact with [plants] and [animals] in manners inconsequential for humans, though prudent humans avoid locations where spirits have clearly affected other beings. Finally, not all spiritual beings are jine. For instance, when a

4 5

Literally, a child of Adam (Bailleul 1996), indicating derivation from Islamic traditions. An Arabic loan word.

71 human dies, his/her hakili (mind, intelligence) becomes the garajik of a newborn. Each human is associated with a garajik spirit, which assists the human in acquiring things of value.6 Acquisition is an important value that helps distinguish [the biospiritual environment] and [the physical environment]. In the Maninka subsistence economy, an individuals ability to acquire valuable things rests on his/her ability to access natural resourcesespecially soil fertility, water, geological materials, microclimate, plants, and animalsand avoid natural hazardssuch as microclimate-induced illness, falling on slopes, and meteorological dangers while also avoiding conflict with powerful beings. [The physical environment] is composed of physical features that indicate the spatiotemporal distribution of natural resources and hazards. However, some features are owned, occupied, or otherwise possessed by powerful beings, especially humans and jine. The concept of possession, which connects physical things to beings, causes individuals to see spiritual and social meaning in physical features, and transforms these into components of [the biospiritual environment]. Components of [the physical environment] are considered neutrally available for use to all people, but those of [the biospiritual environment] are accessible only to people who have socially granted use rights, or who have the spiritual knowledge and power to overcome or appease jine. Subjectivity makes it difficult for informants to divide their world into physical and biospiritual components (Rappaport 1979), but this is clearly practiced when Maninka individuals discuss natural resources. A hunter can describe a location by saying the warthog was past the kna (clearing, field) on the path to the river without suggesting anything about property rights to the field, which is communicated by evoking its social context with a

Bailleul (1996) translates garajike as luck, which is a simplification: Hakara garajike ka nyi means Hakaras garajike is good, but more idiomatically Hakaras luck is good or Hakara is lucky.

72 possessive phrase like nymbi ka furu (Nymbis field). Possessives allow speakers to emphasize resource ownership and thus imply accessibility for each listener according to personal identity and history, moving a discussion from physical features to property rights. Anthropogenic physical features are one type of clue individuals use to locate natural resources, but these features also carry the abstract, socially determined attribute of possession, which limits accessibility to resources associated with these features. Similarly, all artifacts are owned, yet raw materials have no inherent ownership: anyone can use a kuru ge (angular block of sandstone) unless that kuru ge becomes, for instance, buramakan ka si kuru (Buramakans grinding stone). Ownership can lapse for enduring artifacts like abandoned settlements, so that they may become simply physical features. However, due to their past association with humansincluding perhaps subaga (sorcerers) or disguised jine (spirits)these features carry much symbolic meaning and may not be safe to use. The physical environment. [The physical environment] consists of all non-living, physical features of the environment, and comprises three major categories (Figure 3, p. 94). Ala ka baara (the work of Allah) and mg ka baara (the work of humans) clearly differ, but these two, along with [animal-created features], share some subordinate categories. Ala ka baara and mg ka baara are unproductive secondary lexemes (cf. Conklin 1962; Kay 1971); there is no evidence that baara carries a broader meaning equivalent to [the physical environment]. Ala ka baara includes physical features created by ala (Allah), the omnipotent spiritual force. Possession of these features is generally impossible, and thus they remain permanently part of [the physical environment]. Mg ka baara comprises anthropogenic physical features, which may be part of either [the physical environment] or [the biospiritual environment], depending on their ownership status. The noun baara, often translated as manual

73 work or labor (Bailleul 1996), also carries the broader meaning action. Ala ka baara captures both senses, as when informants remarked, humans do not dig caves; they are the work of Allah, or, after a destructive windstorm, the work of Allah was too powerful. In mg ka baara, the sense action pertains to social interactions, while manual work or labor refers to physical features. This latter sense underscores that anthropogenic physical features, like [features created by animals], are manipulations of ala ka baara, and not aboriginal creations. The reference to Allah indicates the historical influence of Islam, not the religiosity of informants. Few Bafing Maninka are Muslim, but Allah designates the omnipotent force in the Maninka belief system (Brun 1907; Tauxier 1927). All spirits are subordinate to Allah, but ala ka baara does not include the actions of subordinate spirits. For example, when asked if instances of garajik ka baara (work of [a] garajik)such as a hunters finding gamewere also ala ka baara, informants said universally these are not the same. Allah creates and maintains the physical environment in which spirits, humans, animals, and plants act. While Allah affects the actions of these beings, such influence reflects Allahs will, and is not Allahs baara (work). [Features created by animals] is of limited conceptual extent. It includes the few enduring features created by animals, such as tun (termite mounds). Abstractly, the animals creating these features possess them, but such possession means little to humans and accessibility to resources held in these features is unrestricted. The components of each of these primary divisions of [the physical environment] are described in the following sub-sections. The work of Allah. Ala ka baara directly includes four categories. Dugu (earth) comprises physical features associated with the ground, including water bodies and certain

74 microclimatic features (specified below). Features composing dugu are above those composing ju kr (the deep subsurface) and below those composing san (the sky). Ju kr7 designates features that are mainly unknowable to humans because of their sub-surface location. San includes all features considered to originate above the ground surface. I use celestial features to designate components of san and terrestrial features for components of dugu. Funteno (temperature) is a usually invisible feature permeating all components of dugu, san, and ju kr. Ju kr is a poorly developed category that some informants divide into dugu ju kr (the deep subsurface of the ground) and ba ju kr (the riverbed) (Figure 3, p. 94). However, for most people ju kr means only the deep subsurface of the ground. Personal history determines an individuals perception of ju kr. Most informants have direct or indirect experience with dugu ju kr via well digging, but few have substantial experience far from Solo, where humans can directly access the bottoms of all water bodies. Informants with experience on the Bafing River, though, consider the riverbed as unknowable as the deep subsurface of the ground. Funteno is a polysemous term that also designates the conceptually most salient temperature state, funteno (hotness), as opposed to nn (cold) and sumaya (coolness) (Figure 3, p. 94). Temperature permeates all physical features, and changes predictably due to interactions of certain celestial and terrestrial features. Tilo (the sun) especially influences temperature changes, but other features have important effects on microclimate. Funteno (hot

Ju means base, foundation, or below/behind [a thing]. Ju kr is: 1) an adjectival phrase (underneath the base [of a thing]), and, in the sense of focus here, 2) a compound noun taking a postposition (e.g. subo te sr ju kr la (mammals are not found in the deep subsurface). A postposition is grammatically identical to a preposition, but follows a noun.

75 temperature) and nn (cold) can be dangerous, especially if magnified by other features. Thus, farmers do not clear all trees from fields, although this would allow greater crop plant density, because nining (shade from trees) is the most important form of sumaya (coolness) moderating midday heat. Other types of heat indicate poor sites for farming or settlement. Although solar heating may cause dugu funteno, its intensity is mainly due to ground surface characteristics: it occurs at night or in shade where nyama (soil healthfulness) is poor. Similarly, not all shade brings sumaya: coolness persists all day in sites with good soil and low spirit activity, but bad sites remain hot even if shaded. San (the sky) is a well-developed category whose subdivisions indicate how observed features relate to the spatiotemporal distribution of precipitation and changes in air temperature (Figure 4, p. 95). San comprises funio (air), kuro (haze), kabo (clouds), san (weather), tilo (the sun), kalo (the moon), lolo (stars), and klbomboli, the locally dominant, northeasterly storm track. San most frequent means weather. Several types of weather are recognized, all associated with rainfall or potential rainfall. The most salient weather is san ji (rain), often called just san. Weather is perhaps the most discussed feature of the biophysical environment, due to its obvious importance in farming. Other celestial features differ according to how they relate to precipitation and microclimatic change. Types of funio (air, wind) and kabo (clouds) interact with other celestial features to produce weather, while these and kuro (haze) affect terrestrial microclimate. For instance, munkun (fog), a type of cloud, can cause dangerously cold air temperature, while kuro (haze) intensifies hot air temperature. Both situations can cause illness for humans, livestock, and crops, depending on site soil and topography.

76 Tilo (sun), kalo (moon), and lolo (stars) are monotypic categories that indicate temporal change. In the past, they may have carried spiritual meaning (cf. de Ganay 1949; Tauxier 1927; Zahan 1950), but currently they mark time without having strongly expressed meanings. Sumaya ni tilimo naaningo8 (the Milky Way) also indicates seasonal change, but is considered a type of kuro (haze). The most developed category subsumed in ala ka baara is dugu (earth), which shares some subordinate categories with mg ka baara. Dugu directly subsumes six categories, in which physical features are differentiated based on topography, hydrology, ground characteristics, and vegetation, each of which embody a set of natural resources and hazards. Additionally, synthetic assessment of all site characteristics represents another criterion by which features are classified. Informants classify [vegetation] by structure or composition (Figure 5, p. 96). The covert category [compositional vegetation] potentially includes many subordinate categories because these are distinguished according to the most salient species in a site (cf. Sow & Anderson 1996), and dozens of species are salient and locally abundant (Duvall 2001). In practice, few compositional vegetation types are recognized, either field vegetation or stands of economically important wild plants (Figure 4, p. 95). [Structural vegetation] types are either tu (vegetation with high stem density and high stature) or kna ge (vegetation with low stature); people can see long distances in kna ge, but not in tu. Short grasses dominate kna ge, which is characteristic of many, but not all, kna (clearings), one of several [land-cover] types discussed below. Different types of tu have high densities of trees, bamboo, or tall grasses.

Literally, the boundary between sumaya (the wet season) and tilimo (the sunny season).

77 Topography is classified in the covert taxon [land forms], which includes wu (cavities), [depressions], and [elevations]. Several criteria differentiate topographic features. First, many features differ according to surface drainage, especially types of [depression] (Figure 6, p. 97). Many depressions are types of ji jigi silo (drainage channel), distinguished mainly by sideslope form. Large, bowl-shaped depressions are distinguished based on drainage network: a solon contains several creeks, a kubo one, and a dinga none. Second, the effects of topography on microclimate also differentiate features. Several features have characteristic degrees of shading, such as types of wu (cavities) (Figure 6, p. 97). Hanhan (caves) contain large, permanently cool areas, and many wrn (pits) retain moisture within narrow openings. Degree of shading also distinguishes some types of [depression]. Deep features like kun sa (drainage channel head) and gouga (gorge) are frequently shaded, while shallower features like bilan da (drainage channel mouth) and hara (swale) are not. Third, slope form differentiates elevated features (Figure 7, p. 98). The concept slope is covert to many older people and most women, but many younger men, who have more exposure to French through labor migration and radio, label this concept koti, from cte (French: slope). Many slope classes are based on how easily they may be climbed: a tinti (rise) is barely noticeable when walking, but a haya (drop) cannot be climbed or descended. Landscape position and substrate also differentiate [elevations]. For instance, rice cultivation is possible on both types of goungou because these are along permanent water bodies; a gongoli (hillock) has the same shape and soil characteristics, but hillocks are not uniformly arable because they occur throughout the landscape. Konko (hill) and kuru (bedrock outcrop) are very salient,

78 differentiated by substrate, not slope form: konko are the edges or remnants of ferricrete crusts, while kuru are dolomite or sandstone outcrops. Permanency, size, and origin distinguish [water bodies] (Figure 8, p. 99). Within the categories of permanent and seasonal, many [water bodies] differ according to the duration water is present during the year or longer periods of time. For example, both gibingibin and ji ja balo are permanent water pools in deep spots in creeks, but a gibingibin is less likely to dry in droughts because it has a rock bottom and occurs in a cavity, not a muddy hole like a ji ja balo. Many water bodies are distinguished by size: a ba (river) is larger than a k ba (creek), and a sakanbe (spring) has more abundant flow than a tondi ji (seep). The origin of water bodies also is important: a kuru bake differs from other flowing water bodies because its water does not belong to a drainage channel, but comes from drainage through soil overlying exposed bedrock. Geologic and soil resources are classified in the category dugukolo9 (ground). There are six components of dugukolo (Figure 9, p. 100). First, ng (organic matter) is surficial and decomposed litter that provides habitat for some animals and enhances the inherent fanga (strength, chemical fertility) of soils. Second, nyama is a gaseous substance that emanates from the ground surfaceespecially from soilthat controls the healthfulness of a parcel of ground. According to an informant, ng and chemical fertilizer are the same; nyama is not the same, but [is] like gaseous pesticide the ground sprays up and makes [some things] sickly even if they grow. Nyama can be good or bad, depending on the site and the being exposed to it. Crops may grow in sites with nyama that is bad for people, but these crops cannot be safely eaten. Third, sumaya (moisture) consists of both nma (soil moisture) and kombo (dew),

Literally, [the] ground[s] bone.

79 which is moisture that has ascended from the ground. Like nyama, sumaya is associated mainly with soil, but types of rock also have varying moisture characteristics. The final three components of dugukolo (ground) are more finely differentiated (Figure 9, p. 100). There are three types of br (gravel), distinguished by particle size. Bogo (soil) is classified based on arability, texture, and color. This category centers on bogo (loam). Bogo and kny (sandy loam) are preferred for farming; less preferred and non-arable soils are clayey or silty. This division reflects the demands of local staple crops: millet prefers welldrained soils, while peanuts cannot be easily dug from dry, fine-textured soil. Kuru (rock, stone) is classified according to which aspecthardness, form, use value, or landscape locationis most salient. Five types of rock are identified by hardness, four by form, three by use value, and three by landscape location (Figure 9, p. 100). Of these taxa, only nari kuru and kaba kuru, both identified by hardness, correspond to rocks recognized by geologists: dolomite and sandstones of the Manantali series, respectively. Clusters of features, not any single characteristic, differentiate types of kuru (cf. Hunn 1976). Many salient features co-occur because these are inherently related, such as how hardness leads to the typical shape of particular rock types. Nonetheless, one feature is considered most salient for each type of kuru, even if the mutual predictability of this and another feature means that the second is, objectively, as characteristic as the first and explicitly recognized as such. For instance, kuru ge (white stone) and kuru fing (black stone) are classifications based on form, although, as their names suggest, their colors are also distinctive. The only stones fitting the size and shape criteria for kuru ge are made of kaba kuru, light-colored sandstone. Indeed, kaba ge is a synonym for kuru ge. Similarly, the only rock that forms stones the size and shape of kuru fing is nari kuru, relatively dark-colored dolomite. San galima kuru (literally thunderstone) is considered to form where

80 lightning strikes the ground. Archaeologists call these celts, or Neolithic polished-stone axe heads (Davies 1967). Finally, jaman kuru (clear quartz) is apparently derived from diamant (French: diamond). Diamonds are not found locally, but French geologists prospected for them in the early 1900s (Varlet et al. 1977) and the name may derive from this contact. Although these separate classifications of topography, hydrology, vegetation, microclimate, and ground surface features are important, most informants consider them altogether when classifying or describing parts of the landscape. This synthetic view produces a separate classification of [land cover]. Land-cover types are either anthropogenic or a type of dan (non-anthropogenic land cover) (Figure 10, p. 101). Dan is a concept laden with meaning, since jine (spirits) occupy parts of the landscape with non-anthropogenic land cover (Brun 1907; Cashion 1982). Dan is the root of danso (hunter) and dansoko (hunters prowess), indicating that hunting represents mastery of the dan and its spirit occupants; exterminating pest animals in fields is not considered hunting, but field management. Dan is subdivided between land cover for which vegetation, landscape position, or topography are most salient. Many land-cover names come from the names of dominant soil types or topographic features but are distinguished grammatically as locative nouns requiring postpositions in all usages. Land-cover types with such names are not simply soil or topographic classes. For example, both kakakure and kuru ge to have kny soil, which is arable, but neither land-cover type is arable. Soil in a kakakure shallowly overlies bedrock, and thus has poor soil moisture characteristics; kuru ge to sites are arable except for the abundance of the grass ngl (Cenchrus ciliaris), a weed. Other landcover names, such as lemukan (arable woodland with sandy soil), take postpositions only when used as objects. Land-cover types for which vegetation is most salient are either kna

81 (clearings) or [not kna], an unlabeled category. Both categories have multiple subdivisions relating primarily to ground characteristics. All land-cover types associated with landscape position are types of mako (creekside), differentiated on ground and vegetation characteristics. Finally, land cover for which topography is most salient are associated with kuru (outcrops) and konko (hills). Although many of these cover types have names derived from types of slope, they are not topographic classifications but require postpositions in all uses. Thus, kuru sinbe he refers to areas found at a kuru sinbe (outcrop toeslope), which have fertile, deep soil, tree-dominated vegetation, good soil moisture, and many colluvial boulders. These areas are valued for agriculture, but not for settlement due to the risk of rock fall. The work of people. Mg ka baara comprises enduring physical features created by humans. Since the act of creating these features imparts possession to their creators, components of mg ka baara belong primarily to the biospiritual environment until their possession lapses. Nonetheless, the context in which a feature is referred determines whether it is perceived as information about the distribution of natural resources, or as an indication of use rights. Anthropogenic physical features are profoundly dualistic, being always, to some degree, part of the biospiritual and physical environments. Many components of mg ka baara belong to taxonomic categories that also include features that compose part of ala ka baara (Figure 3, p. 94). One major division of mg ka baara is [land cover], shared with ala ka baara (Figure 10, p. 101). Anthropogenic land-cover types are differentiated by use. Use distinguishes settlement sites from agricultural clearings and fallows. Anthropogenic clearingsboth furu (fields) and gaso (unfarmed clearings)are part of the broader land-cover category kna (clearing). Significantly, some types of anthropogenic clearingsuch as millet fieldsdo not

82 have kna ge vegetation. Past use characterizes manyang (fallows). Tree density in fallows varies from grassland to forest, but the criterion of past use lumps fallows into a single category regardless of between-site differences in soil, vegetation, or other features. The other major component of mg ka baara is [artifacts], items or structures whose endurance allows them to outlast knowledge of their ownership and become simply physical features. Short-lived artifactslike baskets, fences, or hutsare inherently part of hadamadnya (humanity) since their ownership is never in question. As one informant said, the belongings of people that [disintegrate] if left [without maintenance] are like [antelope] horns. They are hard and strong, but once the [antelope] dies the horns are soon gone. When a man dies his sons may maintain his hut, but when they go [to live elsewhere] his hut will fall []. [However,] some things people can build dont fall [for so long that] we dont know who made them. There are three classes of artifact[manufactures], [structures], and [works] distinguished by form and use (Figure 11, p. 102). Features made by animals. The covert category [features created by animals] includes only kome (salt licks), tun (termite mounds), and wu (holes) (Figures 3, 6, & 11, pp. 94, 97, & 102). Notably, termite mounds are conceived as a type of digging, and thus are part of the category tun (diggings) that also includes parts of ala ka baara and mg ka baara. Discussion Manding cultural ecology. Many physical geographic terms used in Bafing Maninka occur in other Manding dialects, as expected based on their linguistic similarity. Although published vocabularies are incomplete and published glosses often imprecise, Manding dialects share terms referring to broad conceptual categories of physical features. For instance, equivalents to the Bafing Maninka terms san (sky), kaba (cloud), konko (hill), tinti (rise),

83 ba (river), k ba or k (creek, stream), bugu (farm), tumbun (ruined settlement), kolon (well), and others are reported in many Manding dialects (e.g. Anonymous 1906; Bailleul 1996; Bassett & Koli Bi 2000; Bazin 1965 [1906]; Bernus 1956; Delafosse 1929; Delafosse 1955; Flutre 1957; Gamble 1987; Gregoire 1986; Lem 1948; Spears 1965; Tabor 1993; Travl 1913). Manding physical geographic vocabularies are not entirely uniform, however. Some differences reflect history. Manding peoples expanded throughout West Africa in the 13th-15th centuries (Hodge 1971), encountering speakers of other African languages. From the 11th century, Islamic traders spread Arabic across Sahelian Africa, and from the 1400s European traders spoke several languages along the coast (Oliver 1977). Manding groups have had varying exposures to these and other influences, and have borrowed words that indicate different physical geographic knowledge of non-Manding groups. Perhaps the most widespread loan word is sahelo (north), from Arabic. In the southern portion of the Manding area, where Islamic traders were less active historically (Dalby 1971), sahelo is less frequently used, in favor of the Manding words knyka or kogodugu (Bailleul 1981: 286; Delafosse 1929: 549; Travl 1913: 86). The Bafing Maninka are adopting cte (French: slope) for a concept that appears to have been previously covert (Figure 7, p. 98), while Gambian Mandinka and Guinean Kuranko have borrowed non-Manding words (from Wolof, Jola, and Kissi languages) for features that do not appear to have an equivalent Manding name (Carney 1991; Fairhead & Leach 1996; Gamble 1987). Such loan words may represent past adaptations to previously unknown natural resources or hazards, or instances when Manding groups came to distinguish physical features with increased detail following contact with outside groups (cf. Goodenough 1966).

84 The meaning of some widespread terms varies between Manding groups. Some polysemous terms in Bafing Maninka have fewer meanings in other dialects. Particularly, dugu, meaning earth, village site, and population of a village in southwestern Mali (Figs. 3 & 11, pp. 94 & 102; Cisse 1970; Lem 1948), means only earth for Kuranko speakers in southern Guinea, while so means village (Fairhead & Leach 1996). Other widespread terms do not refer to precisely the same physical feature in all areas, indicating the concept behind a term may be broader than the limited set of physical features it names in a given area. For instance, in southwestern Mali the term tu (forest) refers to vegetation that ranges from wooded grassland to forest, but in southern Guinean Kuranko tu means only forest; yrn denotes woodland and frn, wooded grassland (Fairhead & Leach 1996: 204). The Kuranko use these terms to describe vegetation succession. Since equivalent terms are absent in Bafing Maninka, these groups may not similarly understand successionwhich certainly differs qualitatively between the two areas for ecological reasons. Some terms have more complex internal taxonomies in particular Manding dialects than others. For instance, the Gambian Mandinka term faro (swamp: Carney 1991), equivalent to hara (seasonally flooded grassland), is subdivided into at least four types depending on flooding periodicity (Carney 1991; Gamble 1987). These types of faro seem equivalent to Bafing Maninka [land-cover] types (cf. Carney 1991). The subdivision of faro contrasts with the singularity of hara, and indicates variation in agroecological knowledge between the Mandinka and Maninka, and, ultimately, differences in the biophysical environment. The Gambia is a low-lying coastal nation along the Gambia River, where water levels are affected by precipitation seasonsthe sole influence in hilly southwestern Malias well as by tides (Carney 1996; Carney 1991; Michel 1973). Rice is the staple of Gambian farmers; the different types of

85 faro represent classification of the suitability of flooding zoneswhich have distinct hydrological, soil, slope, and vegetation characteristicsfor different rice varieties. In southwestern Mali, rice is a minor crop and most informants recognize only site-specific variation in agroecological characteristics that makes a hara either suitable or unsuitable for rice. In contrast, Bafing Maninka distinguish many [land-cover] types associated with elevated features (Figure 10, p. 101), corresponding to site suitability for the staple crops millet and peanut. Gambian Mandinka appear to have a less detailed classification of upland areas (Carney 1996; Carney 1991; Gamble 1987; Schroeder 1999; Schroeder & Suryanata 1996). Topography is also less complex in The Gambia (Michel 1973), suggesting there are fewer elevated features the Mandinka could recognize independent of use value. Other African ethnolinguistic groups that occupy upland areas, including the Mende in Sierra Leone (a Manding group), have fairly well developed topographic classifications (Osunade 1987). The incompleteness of published vocabularies precludes strong conclusions, but within a widely shared conceptual framework different Manding groups apparently have distinct taxonomies of locally diverse or highly valued physical features (cf. Tabor 1993: 32). This finding supports other research emphasizing the importance of special-purpose taxonomies in classifications of physical features. Ethnobiological research has shown that humans classify plants and animals in highly predictable ways. General-purpose (or natural) classifications of plants and animals are similar across cultures, suggesting a pan-environmental, general-purpose taxonomy of living things (Berlin 1992; Brown 1984). A pan-environmental taxonomy may exist because local florae and faunae do not represent continua of variation, but comprise discrete groups of morphologically distinctive organisms separated by objective

86 discontinuities in the observed range of variation, suggesting a clear classificatory framework (Hunn 1976; Hunn 1977; Malt 1995). Differences between physical featuressuch as landforms, soils, and vegetationis less discrete (WinklerPrins 1999). There are certainly objective discontinuities in the physical environmentsky/earth, elevation/depression, grassland/forestbut within the framework suggested by these discontinuities there are few obvious breaks in the range of variation exhibited by particular classes of physical feature. The nature of continuous variability increases the importance of classificatory criteria in determining taxonomic structure. Since classificatory criteria for sets of items with continuously variable features are culturally subjective, classifications of physical features may be dominated by special-purpose (or artificial) taxonomies (cf. Ellen 1993). Indeed, Williams and Ortiz-Solorio (1981) show that a Nahuatl/Spanish soil taxonomy from central Mexico differs from that of technical soil science because the folk taxonomy classifies surface soils (important to Nahuatl farmers) rather than soil profiles (important to soil scientists). Similarly, Zimmerer (1996; 2001) describes how Andean farmers in Peru and Bolivia classify landscape units. Some of their landscape units correspond to widely used, general termssuch as valley and hillwhile others derive meaning from use-values in potato farming as practiced in the area. Soil and land cover. Maninka farmers in southwestern Mali have a more detailed classification of land-cover types than soil, vegetation, or other features. Soil (and other specific features) contributes only a portion of the perceived arability of a site, which derives from all the natural resources and hazards present. While certain soil types are favored for agriculture, many sites with favorable soil are non-arable due to agroecological constraints posed by topography, vegetation, or non-soil characteristics of the substrate. Similarly, some sites with less favored

87 soils are arable because the total biophysical environment is suitable for certain crops. Soil characteristics vary within most land-cover types, but not enough for informants to recognize soil-based subcategories. The only land-cover types for which a single substrate is diagnostic are non-arable, but not all non-arable land-cover types are associated with one substrate. The synthetic view of soil, vegetation, slope, hydrology, and microclimate embodied in Maninka land-cover categories provides a highly salient and useful indication of agroecological potential for Maninka farmers. Resource managers should use land-cover terms when communicating with Maninka farmers about land management (cf. Laris 2002; Osunade 1988; Osunade 1987; Pulido & Bocco 2003). The importance of land cover over soil, or any single feature, in classifying site arability is a widespread, underemphasized aspect of local knowledge systems (cf. Denevan & Padoch 1988; Fleck & Harder 2000; Frechione et al. 1989; McGregor 1994; Osunade 1988; Pulido & Bocco 2003; Shepard et al. 2001; Verlinden & Dayot 2005). Many works characterized as describing local soil types actually describe land-cover categories, which often include soil assessment but are not soil types. For instance, Carney (1991: 40) describes how Gambian Mandinka farmers recognize micro-environments based on hydrology, topography, and soil. Soil plays a minor role in distinguishing these micro-environments, yet reviews of ethnopedology consistently categorize Carneys paper as describing local soil knowledge (e.g. Barrera-Bassols & Zinck 2000; WinklerPrins 1999). Land cover and soil are not interchangeable concepts in local knowledge systems, nor are land-cover categories simply a portion of local soil knowledge. Nonetheless, the concepts land and soil are frequently confounded in ethnoscientific publications, suggesting inaccurately that local people do not differentiate soil from some or all other natural resources in a site. For

88 instance, Barrera-Bassols and Zinck (2000: 19) state, there is no clear-cut distinction between soil and land characteristics in local knowledge systems (cf. Barrera-Bassols & Zinck 2003a: 171). They report that topography, land use, and drainage are criteria used to classify soils, without citing specific studies. However, works in their annotated bibliography that apparently support these statements actually do not pertain to local knowledge of soils per se, but of landcover types (e.g. Carney 1991; Kant & Defoer 1996; Osunade 1988). In some primary works, land-cover terms are inaccurately applied to soils found in a given land-cover type: Tabor (1993: 47) translates fouga (= huga [ferricrete hardpan]) as a specific soil type, but this is not a soil term (Fig. 10, Laris 2002). Conversely, some authors refer to soil when actually discussing a broader set of environmental features, comparable to Maninka [land cover] categories. For instance, both Osbahr and Allan (2003) and Osunade (1992) repeatedly state that farmers in their study areas examine land characteristicsi.e. a range of biophysical features, and especially vegetationin determining the suitability of sites for agriculture, but consistently describe this as soil knowledge. While soil characteristics may be an important aspect of site arability, farmers clearly know, and use knowledge, about more than soil in selecting arable sites; these land characteristics could as accurately be described as vegetation knowledge (cf. Fleck & Harder 2000; Verlinden & Dayot 2005). In contrast, in their 2003 paper, Barrera-Bassols and Zinck show clearly how local knowledge may be partitioned to distinguish soil and land cover as separate, though related, aspects of the biophysical environment. They report how Purhpecha farmers in central Mexico conceive land as an integrated whole composed of water, climate, relief, and soils. The Purhpecha classify land according to how its four components interact at a given site; soil is only one of several variables that determine productive potential (BarreraBassols & Zinck 2003b: 237-240). The failure to clearly distinguish soil and land has caused

89 misrepresentation of the conceptual extent and distinctness of analogous concepts in local knowledge systems. Additionally, past research has misplaced the concept soil within folk taxonomies of physical geographic features. For the Maninka, bogo (soil) is one of several components of dugukolo (the ground); it is an intermediate-level taxonomic category (Figure 9, p. 100). This finding contrasts with Williams and Ortiz-Solorios (1981) widely accepted position that soil is conceptually equivalent to plant or animala kingdom in folk taxonomical terms. Linguistic evidence poorly supports this position. Kingdoms are generally unlabelled (Berlin 1992). Thus, several authors have considered soil an exception to this principle since soil is a well-defined, labeled category in all studied languages. More parsimoniously, the fact that soil is labeled suggests it is not a kingdom. Indeed, in most folk soil taxonomies (see citations in Barrera-Bassols & Zinck 2000), soil is a primary lexeme that subsumes categories labeled mainly by other primary lexemessuch as loamthat in turn subsume categories denoted by secondary lexemessuch as sandy loam. In such cases, soil fits only the linguistic criteria for life form, not kingdom (Berlin 1992). Researchers who have misclassified soil may not have studied broad enough samples of pertinent local knowledge systems to observe the concepts full context. Some sources suggest, as shown here for the Maninka, that soil is included in a broader conceptual category that also includes, at the minimum, stone (e.g. Kant & Defoer 1996; Romig et al. 1995; Ryder 1994; Sandor & Furbee 1996). For example, the Purhpecha in central Mexico recognize numerous types of soil using secondary lexemes derived from the primary lexeme echeri (soil), and four types of stone using secondary lexemes based on the primary lexeme tzacapu (stone) (Barrera-Bassols & Zinck 2003b: 239). The authors list both tzacapu and echeri under the

90 heading soil terms, suggesting these are taxonomically equivalent categories subsumed in a category equivalent to the ground. Knowledge of soils in relation to agricultural practice, an important research topic, must not be divorced from broader knowledge of natural resources farmers use in assessing arability. Soil is but one ground-surface feature farmers assess, and the ground surface is but one of several broad classes of physical feature that compose agroecological potential. This discussion of the taxonomic rank of soil in local knowledge systems suggests a broader question about the conceptual scale of folk taxonomic research. Research on ethnobiological classification has indicated that humans universally recognize, if covertly, the concepts plant and animal (Berlin 1992; Brown 1984). These categories exist as natural realms in local knowledge systems at the taxonomic rank of kingdom, the category that subsumes all related objects. How are kingdoms related? In the Maninka conceptualization of the biophysical environment, plant and animal kingdoms are included in a broader category, [the biospiritual environment], which includes all beings and thus contrasts with inanimate features of [the physical environment], even though the two categories are not absolutely separable (Figure 2, p. 93). This higher taxonomic rank is unnamed, because few researchers have considered its existence (cf. Rappaport 1979). Inattention to broad taxonomic contexts has led researchers to misinterpret the taxonomic rank of soil, for instance, and greater attention should be given the overarching structures of local knowledge systems in order to clarify the epistemology of aspects of local knowledge and their scientific analogues. Better knowledge of how different cultures identify and classify physical geographic features will improve understanding of the conceptual foundations of physical geography (Atran 1990; Blaut 1979).

91 Figures for Chapter Two



ala ka baara ju kr ('deep subsurface') san, san hutuma ('sky') {Fig. 6} funteno ('temperature') dugu ('earth') funteno ('hot temperature') nn ('cold temperature') sumaya ('coolness') [water bodies] {Fig. 10} [substrate] {Figs. 11, 12} [vegetation] {Fig. 4} [landforms] {Figs. 8, 9} siya ('lair') nyaga ('nest') kome ('salt lick') [land cover] {Figs. 13} [artifacts] {Figs. 14} dugu ju kr ('deep subsurface of the ground') ba ju kr ('riverbed') kuru funteno ('humidity over damp rock') bin funteno ('humidity over damp grass') dugu funteno ('heat of the ground') nining ('shade under trees') siniware ('shade from clouds')

[features made by animals]

mg ka baara

Figure 3. Main categories of the Maninka physical environment. Line formatting indicates taxonomy for the three subdivisions of [the physical environment]: solid lines= ala ka baara; dotted lines=[features made by animals]; dashed lines=mg ka baara. Categories without shading belong uniquely and totally to ala ka baara; lightly shaded categories belong uniquely and totally to [features made by animals]; darkly shaded categories belong uniquely and totally to mg ka baara; categories outlined in gray subsume categories that include features belonging to more than one of the three primary subdivisions. Commas separate synonymous terms. For categories followed by braces, internal taxonomies are shown in the figures indicated. Intersection of mg ka baara and [features made by animals] with the [biospiritual environment] is not represented here; see Fig. 2 and the text for description of these intersections.









103 Chapter Four: Human and environmental causes of floristic patterns in southwestern Mali

Abstract This paper presents the results of vegetation studies conducted in southwestern Mali, which lies in the semi-arid Sudanian bioclimatic zone. The dominant view of the Sudanian zone is that vegetation distribution and composition has been heavily affected by cultivation, and that anthropogenic impacts create distinct floristic patterns. Vegetation sampling occurred at 217 sites with disturbance histories determined through interviews of local residents. For each site, data was collected on edaphic characteristics, topography, hydrography, disturbance, and (for previously cleared sites) time since abandonment. Floristic analyses (hierarchical cluster analysis, indicator species analysis, multi-response permutation procedures, and non-metric multi-dimensional scaling) revealed that: 1) Fifteen vegetation types are recognizable, with different levels of floristic distinctness. 2) Most vegetation types are reliably indicated by various species, and are linked to specific environmental or disturbance conditions. 4) Edaphic features, especially bedrock characteristics, and landscape position explain most floristic variation. The importance of particular sandstone bedrock conditions in the maintenance of mesophytic refugia is underscored. 5) Finally, three different vegetation responses to human disturbance are apparentvegetation turnover, homogenization, and resiliencedepending on site-specific biophysical and socioeconomic conditions. In particular, vegetation in abandoned settlements and abandoned fields show distinct response to disturbance, because these two human activities differ in terms of ecological impact. However, effects of human disturbance are difficult to identify due to the absence of undisturbed vegetation in arable sites. Keywords

104 Vegetation; biogeography; ecology; Mali; agriculture; disturbance; NMS; MRPP; ISA

Introduction In the semi-arid tropics, variation in woody vegetation structure and composition corresponds almost entirely to variation in edaphic and hydrological features (Bourlire 1983; Breman & Kessler 1995; Bullock et al. 1995; Cole 1986; Furley et al. 1992; Lawesson 1995; Lawson 1986; Meave & Kellman 1994; Meave et al. 1991; Schnell 1971; Scholes & Walker 1993; White 1965; White 1983). Woodlands and dry forests dominate semi-arid tropical regions, particularly in sites with relatively deep, arable soil. However, these regions also host forests, grasslands, and bushlands in locations predictable from soil and hydrological characteristics. For instance, in Sudanian West Africa forest patches enriched with mesophytic, Guinean species occur in sites with elevated soil moisture content, while grasslands and bushlands host xerophytic Sahelian and Saharan plants in sites with shallow or infertile soil (White 1965). Adopting a regional scale of observation in the semi-arid tropics underscores the importance of edaphic and hydrological factors as sources of variation in vegetation composition and structure. A less expansive scale of observation that focuses on landscapes (areas of hundreds of square kilometers) increases the significance of human activities as a source of vegetation variation. While constraints imposed by edaphic and hydrological variation remain evident in many landscapes as riparian forest corridors and vegetation patches associated with unusual substrates (Breman & Kessler 1995; Bullock et al. 1995; Lawesson 1995; Meave & Kellman 1994; Meave et al. 1991), the structure and composition of most vegetation types have been profoundly modified through human management.

105 Human activities are especially significant as a source of vegetation variation in semi-arid Africa because of the continents extremely long human history. In particular, the composition of African woodlands reflects millennia of human disturbance via fire, settlement, agriculture, and edible fruit gathering (O'Brien & Peters 1998; White 1983). Humans have modified the local, regional, and continental distributions of valuable, indigenous fruit trees, and increased the absolute and relative abundance of these species around settlements (Boffa 1999; Chevalier 1947; Duvall 2006; Maranz & Wiesman 2003; McGregor 1994; Pullan 1974). More significantly, vegetation clearing for agriculture and the use of fire in farming, animal husbandry, and resource management have probably resulted in increases in the abundance and distribution of species adapted to these disturbances (Bassett & Boutrais 2000; Belsky 1995; Belsky et al. 1993; Breman & Kessler 1995; Laris 2002; Nyerges 1989). Intentional or unintentional alteration of the distribution and abundance of certain species may create anthropogenic forest (Amanor 1994; Avenard et al. 1974; Fairhead & Leach 1996; Spichiger & Pamard 1973) and bushland (Audru 1977; Bassett & Boutrais 2000; Bassett & Koli Bi 2000) patches in some semiarid areas. Human activities are clearly significant in vegetation variation as viewed at a landscape scale. Indeed, botanists and vegetation scientists working in West Africa have historically seen woody vegetation characteristics as profoundly indicative of past human activities (e.g. Aubrville 1949a; Chevalier 1933; Schnell 1976), and have thus overlooked biophysical sources of vegetation variation while overemphasizing possible human causes (Cole 1986). This interpretation of vegetation characteristics has been frequently coupled with the view that woody vegetation in semi-arid West Africa is degraded relative to its supposed, aboriginal condition, and has been expressed in terms that assign blame for the purported degradation on the supposed

106 destructiveness of indigenous resource management (Bassett & Crummey 2003; Duvall 2003; Fairhead & Leach 1998; Leach & Mearns 1996; Richards 1985; Turner 1993). Of course, many researchers have avoided overtly political statements about indigenous practices, but have sought only natural vegetation, ignoring plant communities considered disturbed or otherwise not representative of natural biophysical influences. The results of such studies have underscored the significance of edaphic and hydrological factors in vegetation variation (e.g. Adam 1956; Lawesson 1995; Nasi & Sabatier 1988; Roberty 1940; White 1965; White 1983). Partially in response to approaches to vegetation study that represent humans as an exogenous, unnatural influence on vegetation, students of cultural ecology have focused on vegetation in fallows, fields, pastures, and/or settlements. Such studies have shown that indigenous farming, animal husbandry, and fire management are not inherently destructive of vegetation, and that anthropogenic vegetation change is not necessarily, or even frequently, the deforestation often assumed (e.g. Amanor 1994; Bassett & Koli Bi 2000; Chidumayo 2004; Devineau 2005; Devineau 2001; Fairhead & Leach 1996; Lykke 1998; McGregor 1994; Nyerges 1989). Nonetheless, few studies in any field of inquiry have simultaneously considered both human and biophysical causes of vegetation variation in Sudanian West Africa (e.g. Breman & Ciss 1977; Breman & Kessler 1995), even though these causes interact to create floristic patterns in populated landscapes (Berkes & Folke 1998; McDonnell & Pickett 1993; Vale 1982). Most studies of floristic variation in semi-arid West Africa have privileged either human activities or biophysical variation as explanatory factors, regardless of scale of observation or views about the destructiveness of human activities. An emerging challenge in natural resource management is to develop land-use ecologies that account for vegetation change without resorting to oversimplified or inappropriate

107 explanatory frameworks (Turner 2000). Long-term conservation and management of biodiversity resources depend on our ability to situate human activities within ecosystem processes (Baker 1992; Bellemare et al. 2002; Berkes & Folke 1998; Micheli et al. 2001). Humans have been integral components of African ecosystems for millennia. Through much of the 1900s, many natural resource scientists recognized the extremely long history of resource management by African farmers and herders by blaming them for supposed environmental destruction. Dominant narratives of widespread, anthropogenic deforestation in West Africa have proven inaccurate or unsupportable (Bassett & Koli Bi 2000; Duvall 2003; Fairhead & Leach 1996; Ribot 1999). Nonetheless, indigenous agricultural and settlement practices do alter vegetation structure and composition (Boffa 1999; Lykke 1998; Maranz & Wiesman 2003; Nyerges 1989; Nyerges & Green 2000; Schreckenberg 1999). Anthropogenic vegetation changes overlie a pattern of edaphic and hydrological constraints to vegetation development, which altogether structure semi-arid African ecosystems. How do anthropogenic vegetation changes interact with floristic patterns that exist independently of humans? Most studies of the effects of human activities on woody vegetation in West Africa have been limited to specific portions of landscapesareas with relatively deep, arable soil (e.g. Bassett & Koli Bi 2000; Devineau 2005; Fairhead & Leach 1996; Lykke 1998; Schreckenberg 1999)where human impacts are greatest and most evident. Indeed, these studies have sought to identify and describe anthropogenic vegetation characteristics, often in order to contribute to resource management debates. However, recognizing relationships between human activities and ecosystem structure requires more thorough vegetation study; the significance of anthropogenic vegetation change depends upon the full range of vegetation variation in focal landscapes (Baker 1992; Vale 1982). Focusing on human origins of vegetation

108 characteristics, on the intentionality of anthropogenic changes, or on their perceived benefit/detriment to preferred uses (or meanings) of focal landscapes risks the conceptual excision of humans from the biophysical system of study and the privileging of human activities over non-human factors in explaining observed variation (cf. Craghan 2004). While human intentions, perceptions, and goals are vitally important in the context of resource management and conservation, these are less important in understanding the physical processes that spatially structure ecosystems, even if some of these processes proceed from human activities. The purpose of the present paper is to examine the relative significance of various biophysical factors, including past human activities, to floristic variation in a southwestern Malian landscape. Although this paper identifies vegetation changes attributable to human activities, its goal is not to identify such changes per se, but instead to identify biophysical factors, including human activities, that may explain observed variation in the composition of plant communities. By considering humans as one of several possible sources of vegetation variation, this paper contributes significantly to identifying and understanding the processes that create of landscape-scale floristic variation and ecosystem structure in semi-arid Africa. Research location Research was conducted in an area of 183 km2 around Solo village, in Malis Bafing Biosphere Reserve (BBR) (Figure 1, p. 150). This area is part of the Manding Plateau, a range of sandstone plateaus in southwestern Mali, that lies in the Sudanian bioclimatic zone. Topographic complexity creates a wide range of habitats in this area. Sandstone plateaus dominated the landscape, rising 200-300 m above surrounding lowlands (IGM 2001). This upper planation surface erodes to form narrow ravines, rocky slopes, and plains with relatively infertile sandy and silty soils. Ferricrete hardpans and bare sandstone surfaces, with very dry

109 microclimates, are common (Dames & Moore 1992; Jaeger & Jarovoy 1952; Michel 1973). Groundwater seeps to the surface permanently or seasonally in some locations where sedimentary layers in the sandstone have been exposed; if seepage occurs in topographically sheltered locations (such as ravines), very humid microclimates exist (Duvall 2001). Elsewhere, permanently moist habitats are uncommon. Soils in most locations are driest from March to June, when average temperature and evapotranspiration peaks (FAO 1984). Precipitation is highly seasonal (June-September) and averages about 1200 mm per year, with high interannual variation (FAO 1984; Leroux 2001). Since the early 1900s, vegetation in the Manding Plateau has been considered prototypical for the Sudanian region (Aubrville 1950; Chevalier 1938; Chevalier 1900). Woodland vegetation dominates flat to sloped areas with good drainage and relatively deep soil, and exhibits weak floristic variation associated with soil and topography (Lawesson 1995; Nasi & Sabatier 1988; Projet Inventaire 1990). Three types of forest may be present in patches at sites with high soil moisture content (Adam 1960; Duvall 2001; Lawesson 1995; Nasi & Sabatier 1988): riparian gallery forests, along permanent waterways, with Raphia sudanica and bamboo (Oxytenanthera abyssinica) characteristic; non-riparian gallery forests, in humid microhabitats on bedrock outcrops, with many Guinean species and often dominated by the tree Gilletiodendron glandulosum, endemic to the Manding Plateau; and non-gallery forests, often on toeslopes with deep soil, with Anogeissus leiocarpus common.

110 Rupicolous bushland occurs in very rocky sites on plateau tops and has a low density of woody plants (Raynal & Raynal 1961; White 1983). Another bushland community, with higher stem density, occupies eroded areas with clayey soil (Nasi & Sabatier 1988). Edaphic grasslands and wooded grasslands occur where a shallow soil layer covers ferricrete or bedrock, and in seasonally flooded swales (Lawesson 1995). Finally, past researchers have considered vegetation in occupied settlements and active fields a single, undifferentiated vegetation type, but have provided no floristic data (Nasi & Sabatier 1988; Projet Inventaire 1990). Abandoned settlements and fields have not been recognized as having distinct vegetation. Evidence of anthropogenic disturbance in many parts of the landscape is not readily apparent (Duvall 2001). All sites, including those labeled undisturbed below, are subject to at least low intensity or frequency disturbance (cf. Vale 2002). The indigenous Maninka people settle and cultivate lowland sites with arable soils and good drainage. Rocky areas, plateau tops, sites with poor soil or drainage, and edaphic grasslands are used only for seasonal livestock grazing, wild plant and honey collection, and hunting (Duvall 2001; Samak et al. 1987). Around settlements, farmers annually burn most grasslands and some woodlands to prevent destructive fires and to prepare fields (Laris 2002). During field clearing and subsequent management, farmers preserve individuals of several tree species with edible fruits (Koenig & Diarra 1998; Samak et al. 1987), which may dominate vegetation for several decades after fallowing. Abandoned settlement sites are recognizable as patches of trees with edible fruits especially baobab (Adansonia digitata) (see Chapter 5). Although Fairhead and Leach (1996) describe a teleological process in which Maninka farmers in Guinea purposefully transform vegetation by planting valuable wild species around settlements, similar practices are not readily evident in southwestern Mali.

111 Farming is seasonal and rain-fed. Immediately around settlements, farmers maintain small maize fields that are farmed each year, but most fields, planted with the primary staples millet, sorghum, fonio, and peanuts, are located farther from settlements and cultivated <10 years before fallowing >10 years (Samak et al. 1987). Arable soil is patchily distributed across the landscape (PIRT 1983), and farmland is limited around Solo. As a result, many farmers must improve their access to farmland by establishing farming hamlets some distance from Solo, but within Solos area of traditional usufruct (see Chapter 2). Most hamlets are occupied only during the farming season and only for relatively short time periodsin most cases, <20-30 years before abandonment (see Chapter 2). Hamlets are usually occupied by only a small number of related, nuclear families who return to Solo after a hamlet is abandoned, and often establish other hamlets after some time in Solo. The practices of hamlet establishment and abandonment represent a shifting settlement system (cf. Stone 1996). Hamlet farming has probably been practiced in the research area for at least several hundred years, but has become increasingly important over the last century (see Chapter 2). Conservationists working in Mali consider hamlet farming a spatially uniform and destructive threat to natural habitat in the area (see Chapter 2), but the cultural ecology of hamlet farming has not been studied. Social changes have led to an increase in hamlet farming elsewhere in southwestern Mali (Koenig & Diarra 1998), but around Solo hamlet establishment has declined following the promulgation of policies to forcibly remove unauthorized settlements from the BBR. In most cases, hamlet establishment is illegal, since national laws prohibit clearing vegetation that has not been cleared for more than 10 years (Prsidence de la Rpublique du Mali 1995). Farmers know this law: Solos residents did not agree to allow

112 vegetation sampling in currently occupied settlements or fields, but were supportive of and assisted in sampling abandoned sites. Data collection This research is based primarily on vegetation sampling, complemented by ethnographic study. Vegetation sampling design had two distinct components: a) a study of chronosequences of abandoned settlement and abandoned field sites, and b) a study of floristic variation across all parts of the focal landscape. The results of these studies together allowed identification of vegetation responses to human disturbance, and the significance of these responses in altering floristic patterns that exist independently of disturbance caused by settlement or cultivation. Sampling site selection and characterization. From January to December, 2004, vegetation sampling occurred at 217 sites. Sampling sites were selected based on substrate texture, bedrock geology, slope, hydrology, and past human use. These five factors are collectively called environmental factors. Additionally, site types were informally defined as locations with similar environmental factors. These informal site types were used to facilitate the selection of sample sites. At the beginning of research, when vegetation sampling began, as wide a variety of site types were sought in order to estimate their range and variety in the research area. After several weeks, when few new site types were being found, research effort focused on sampling: a) an approximately equal number of sites per type; b) widely dispersed examples of each site type; and c) approximately equal numbers of sites either with evidence of past human settlement or cultivation, or without evidence of such past use. For some site types, few sites were sampled because type-defining characteristics were rare. Grasslands were not sampled. The location of each site was determined using a Garmin GPS-12XL unit, and represented as a point corresponding to the approximate center of the area sampled.

113 For each site, the following environmental factors were recorded: Substrate texture: For most sites substrate texture was recorded as a soil texture category (e.g. sandy loam, silty clay loam), determined manually (Midwest Geosciences Group 2003). For sites where 25% of the surface was covered by stones or ferricrete nodules >5 cm in diameter, substrate texture was based on visual estimation of the percent surface area covered by different grain sizes, or consolidated bedrock. Bedrock type: For sites with surficial rock, rock was identified following Varlet et al.s (1977) and Groupement Manantalis (1979) descriptions; determination of subsurface geology followed DNGM (1992). Four types of bedrock were observed: plites (fine-grained sedimentary rocks), Sandstone 0, Sandstone B, and dolomite (using Groupement Manantalis terminology). Ferricrete was also included as a rock type. Further description of geology and geomorphology are included in the discussion section, below, where it is relevant for understanding results. Slope: Slope angle measurements made with an inclinometer over a distance of c.100 m in the approximate center of the area sampled. No sites were sampled in which slope varied by >5. These data were used in their original, quantitative form, or as categorical data reflecting ranges of values (e.g. 5, >10). Hydrography: These categories were based on: the topographic position of a site relative to adjacent parts of the landscape; distance between sample plots and drainage channels; and the presence of permanent water sources directly upslope or adjacent to sample sites. Past use, as described in the following paragraph.

114 All categories for each of these environmental factors are listed in Table 1 (p. 160), which also shows the number of sites sampled per category. All sites with evidence of past settlement (e.g. remains of huts) or cultivation (e.g. rock piles, rock lines, girdled stumps) were classified as disturbed, while undisturbed sites were those without evidence of past use. For each disturbed site, three types of data revealed past use (settlement or cultivation) and time since abandonment: Interviews of past occupants, their relatives, and/or their descendents were used to estimate abandonment date and identify site use (settlement or cultivation). Abandonment dates were estimated by correlating informant life history markers, changes in site occupation status, and datable events, such as national elections. Multiple informants were interviewed for each site, to triangulate date estimates and increase precision (Flowerdew & Martin 1997). For some settlement sites, past occupation was dated from historical documents (Anonymous 1958; de Lannoy de Bissy 1882; Park 1954 [1815]; Projet Inventaire 1990) or aerial photos taken in 1952. These photos also allowed dating use of some abandoned field sites. For three abandoned field sites, the presence of rock piles, rock lines, and girdled stumps attested past use, although only generalized oral historical evidence supported this. If a disturbed site had been occupied at two separate periods, only the most recent abandonment date was used in analyses. Sites were assigned to 10-year age classes; all settlement sites abandoned >40 years ago (y.a.), and fallows abandoned >30 y.a., were grouped together. For

115 some analyses, past-use categories were lumped together based on disturbance type (settlement or cultivation) and time since site abandonment. A concurrent census of settlement sites in the research area revealed 7 occupied and 80 abandoned settlements (see Chapter 2). All abandoned settlement sites (henceforth ruins) that had not been subsequently cultivated (n=50) were sampled. Sampling also included 61 abandoned field sites (henceforth fallows), none of which included ruins. These 111 disturbed sites nearly equaled the number of undisturbed sites (n=106). Vegetation sampling. At each site, ten 250m plots (0.1 ha total area) were established (one site had thirteen plots) following Alwyn Gentrys methods (cf. Phillips & Miller 2002). All woody plants, bamboo, and palms rooted in these plots and meeting size requirements were sampled. Specific data varied by growth form: for single-stemmed trees and shrubs 2.5 cm diameter at breast height (DBH), DBH was measured with a diameter tape and recorded; for multi-stemmed trees and shrubs with at least one stem 2.5 cm DBH, DBH of the largest stem was measured and recorded; for lianas with stems 2.5 cm diameter at any visible point, maximum stem diameter was measured or estimated and recorded; for bamboo, the number of live stems 2.5 cm DBH was counted and multiplied by the estimated average DBH of stems 2.5 cm DBH (usually 2.5-3.0 cm); and for palms having stems (not solely leaves) 1.4 m high, basal diameter of the stem, not including petiole bases, was estimated. Voucher specimens, deposited at the Missouri Botanical Garden (MO), were collected for most species encountered. Additionally, qualitative notes on microhabitat characteristics were

116 recorded for each species. Notes included: specific locations where individuals occurred within patches, relative to geomorphic, geologic, and topographic features; species that occurred in the same or similar locations; and phenological status of each individual observed in samples. At each site, sampling began at a randomly established point; subsequent plots usually began at the end point of the previous plot. The long axis of each plot was oriented in a compass direction different from those of adjacent plots, although in riparian corridors, plots were nearly parallel. At least 10 m separated near-parallel plots. No plots overlapped or included more than one site type. Plot length was determined using a hip chain, while width was estimated from the researchers reach, measured at 1.96 m. At each site, plots were dispersed in an area 200-500 m 10-100 m. Data analysis Summary statistics. For each site, the following summary calculations were made: number of individuals, species, and families, number of individuals per species and family, and number of species per family; density (individuals per unit area) and basal dominance (summed basal areas of all individuals, based on DBH) per species and for all species; diversity, using the Berger-Parker index, which is equal to the proportional abundance of the most abundant species in a sample, and most strongly reflects species evenness, rather than richness (Southwood & Henderson 2000). These summary statistics were used in subsequent analyses, described in the following paragraphs. Floristic analyses. Three complementary analyses were conducted using PC-ORD software (McCune & Mefford 1999): 1) hierarchical cluster analysis interpreted via indicator

117 species analysis (ISA); 2) multi-response permutation procedures (MRPP) of clusters and sites grouped by environmental factor; and 3) non-metric multi-dimensional scaling (NMS). In all analyses, between-site distances were calculated using the Srensen (Bray-Curtis) index (McCune & Grace 2002; Southwood & Henderson 2000). First, cluster analysis was used to identify groups of sites with similar vegetation composition. This classification method has a long history of use in ecology, and many technical descriptions and research applications have been published (cf. McCune & Grace 2002). For the present research, the steps performed by PC-ORD in cluster analysis were: 1) a between-site dissimilarity matrix was calculated from proportional abundance values per site; 2) the sites with the lowest dissimilarity value were linked using the flexible beta method (=-0.25); 3) the information lost by creating this new group (i.e. dendrogram scaling) was calculated using Wisharts objective function; 4) a new dissimilarity matrix was calculated using the new group; and 5) steps 2-4 were repeated until all sites had been grouped together. Linkages and associated objective function values were represented graphically as a dendrogram. Dendrogram pruning was based on ISA, MRPP, and subjective interpretation of the ecological meaning of clusters. Described by Dufrne and Legendre (1997), ISA assesses the fidelity of species to predefined groups of sample sites (i.e. clusters identified through cluster analysis). The indicator value (IV) for each species per cluster ranges from 0% (no indication) to 100% (perfect indication). Interpreting IVs is easiest when done in comparison with statistics on within-cluster homogeneity, such as mean distance. Species with high IVs for clusters with low

118 within-cluster homogeneity are generalists, and IVs for these species decrease as within-cluster homogeneity increases (Devineau 2005). In contrast, IVs for specialist species increase as within-cluster homogeneity increases (Devineau 2005). However, if the number of sites per cluster decreases too greatly, IVs are less indicative of generalist or specialist adaptation, because cluster characteristics become indicative only of conditions in a small number of sites (Devineau 2005). Thus, IVs are useful in identifying the smallest ecologically meaningful clusters identified through cluster analysis (McCune & Grace 2002). For the present research, the steps performed by PC-ORD in ISA were: 1) the proportional abundance (the proportion of all individuals that belong to a species) of each species in each cluster was calculated relative to its abundance in all clusters; 2) the proportional frequency (the proportion of sample sites in which a species occurs) of each species in each cluster was calculated; and 3) these two proportions were multiplied then expressed as a percentage, for each species in each cluster. These percentages are the IVs per species per cluster. The statistical significance of IVs was assessed by randomly reassigning species to groups 1000 times, then calculating IVs for these random reassignments (McCune & Grace 2002). The 22 most informative clusters were subject to ISA. IVs provided no clear indication of which clusters should be retained after pruning. To help determine the optimal set of clusters to retain, MRPP was used to test the statistical significance of the between-group heterogeneity and within-group homogeneity exhibited by different sets of clusters. MRPP is a multivariate, nonparametric method of testing the hypothesis of no difference in species composition between predefined groups of sample units (e.g. clusters identified through cluster analysis). MRPP has been used regularly in community

119 ecology, especially to assess the influence of disturbance and environmental factors on vegetation composition (cf. McCune & Grace 2002; Mielke & Berry 2001). Technical decriptions of MRPP include Biondini et al. (1985), Zimmerman et al. (1985), Mielke and Berry (2001), and McCune and Grace (2002). MRPP essentially assesses the likelihood that observed within-cluster homogeneity and between-cluster heterogeneity for predefined clusters are due to chance, based on randomization of group membership (Mielke & Berry 2001). Specific procedures used by PC-ORD in MRPP analysis are described by McCune and Mefford (1999). In the present analysis, distance measures were rank-transformed, which increases sensitivity as community heterogeneity increases, and makes MRPP results more analogous to those provided by NMS (McCune & Grace 2002). The distance matrix used abundance per species per site; the weighted mean within-group distance () used the standard group-weighting equation, Ci=ni/ni (McCune & Grace 2002). For the present research, the 22 most informative clusters from cluster analysis were subject to MRPP, then the 21 most informative, and so on. Based on interpretation of IVs and MRPP results, fifteen clusters were retained after pruning as the ecologically most meaningful (Table 2, p. 162). Additionally, MRPP was used to assess the significance of these fifteen clusters relative to site groupings based on environmental factors. Finally, NMS was used to assess the relationship between environmental factors, past use, and floristic patterns. Clarke (1993), McCune and Grace (2002), and others describe this ordination method; published applications of NMS in ecology include Kantvilas and Minchin (1989), Tuomisto et al. (1995), and Waichler et al. (2001). NMS was used because several species (especially trees with edible fruit) appeared to have bimodal relationships to disturbance

120 history, being abundant in both ruins and undisturbed forests. The appropriateness of NMS analysis is not limited by nonlinear species-variable relationships (Clarke 1993). Three separate NMS ordinations were undertaken: one each using data from all sites, only undisturbed sites, and only disturbed sites. Ordinations attempted using only sites belonging to the woodland/bushland cluster, described below, were highly unstable, while others using only sites belonging to the forest cluster, described below, were marginally more informative than the undisturbed-site analysis. Analyses proceeded through several iterations testing different combinations of starting configuration, exclusion of rare species, thoroughness settings in PC-ORDs NMS autopilot mode (McCune & Mefford 1999), and number of dimensions. The analyses chosen as final were those with the lowest instability and stress. Specifically, in all analyses, species occurring in less than 5 sites were eliminated, and the medium thoroughness default settings were used (15 runs with real data, 30 with randomized data, 200 maximum iterations, and 0.0001 instability criterion: McCune & Mefford 1999). Final analyses began with the most stable configuration of intermediate trials to avoid local minima. The all-site and undisturbed-site analyses found 2-D solutions, while the disturbed-site analysis identified a stable, 3-D solution. Finally, ordination axes were interpreted by overlying quantitative or categorical variables for environmental factors, including past use. Results Vegetation clusters. Cluster analysis produced a dendrogram in which the primary division splits forest sites from woodland, wooded grassland, and bushland sites (Figure 2, p. 152). Fifteen less inclusive clusters were retained after pruning. MRPP analysis of the two primary clusters shows that they are very distinct compositionally (T=-82.1, A=0.22, p<0.0001; see Table 2, p. 162, for explanation of statistics), which is also suggested by strong IVs for each

121 cluster. The woodland/bushland cluster is strongly indicated by the generalist species Combretum glutinosum and Pterocarpus erinaceus, while the forest cluster is indicated by Spondias mombin, Oxytenanthera abyssinica (bamboo), and the liane Sarcocephalus latifolius (Figure 2, p. 152). Average between-site distance in the two primary clusters is high; less inclusive clusters have lower average between-group distances (Figure 2, p. 152). However, in the woodland/bushland cluster IVs also mostly decrease in less inclusive clusters, while in the forest cluster most IVs increase, indicating that specialist species dominate forest vegetation (Devineau 2005). Several clusters retained after pruning can be lumped together and retain ecological meaning (Figure 2, p. 152). However, MRPP shows that between-group heterogeneity and within-group homogeneity is higher for the 15 clusters than for groupings based on any single environmental factor (Table 2, p. 162). Grouping sites in an intuitive mannerwith disturbed sites grouped by disturbance type, time since disturbance, and substrate texture, and undisturbed sites grouped by vegetation structure and substrate textureproduces the second-best result in terms within-group homogeneity, while the second-greatest between-group heterogeneity is achieved by grouping sites by the binary variable disturbed or undisturbed (Table 2, p. 162). The ecological characteristics of the fifteen clusters retained from cluster analysis are described in the following paragraphs. Summary statistics for these vegetation types are presented in Tables 1 and 3-18 (pp. 160 & 163-178). Clusters 1-3 comprise the forest cluster, and all three consist of sites with moist soil conditions. Clusters 1 and 2 comprise sites with gallery forest vegetation found in moist microhabitats along steeply sloped rock outcrops:

122 Cola cordifolia, Spondias mombin, and Bombax costatum indicate Cluster 1 vegetation. Structurally, no species dominatesthis cluster has the highest diversity of all clusters (Table 3, p. 163)but several are important (Table 4, p. 164). These sites occur on scree slopes, where exposed bedrock occupies 25% of the surface (Table 1, p. 160). Most have permanent springs or lie along seasonal drainage channels. Cluster 2 represents outcrop sites with gallery forest vegetation dominated by Gilletiodendron glandulosum and Hippocratea indica (Table 5, p. 165). Other than these dominant species, vegetation composition is similar to Cluster 1, but compositional heterogeneity is extremely low (Table 3, p. 163). Cluster 2 sites occur in narrow ravines and along cliff tops; in contrast to Cluster 1, all Cluster 2 sites have 25% of the surface covered by bare bedrock, uniquely Sandstone B (Table 1, p. 160). Many component species, especially indicator species, grow in the vertical cracks characteristic of Sandstone B (Jaeger 1950b; Jaeger & Jarovoy 1952; Raynal & Raynal 1961), in which groundwater remains accessible throughout the year (DCTD 1990; Groupement Manantali 1979; Varlet et al. 1977). Bamboo (Oxytenanthera abyssinica) strongly dominates Cluster 3 vegetation, both structurally and compositionally (Table 6, p. 166), resulting in low diversity (Table 3, p. 163). These sites occur mainly in seasonally dry drainage channels (Table 1, p. 160), and their composition includes some species indicative of Clusters 1 and 2 (e.g. Spondias mombin, Saba senegalensis) as well as woodland indicators (e.g. Pterocarpus erinaceus). This cluster includes one fallow site abandoned 0-10 y.a., in which vegetation is structurally not forest.

123 Clusters 4-6 are weakly distinguishable, both individually and as a group (Figure 2, p. 152). Ecologically, these sites are important because Maninka farmers consider most of them non-arable (see Chapter 3). Their soil is mostly silty or clayey, and many overlie ferricrete hardpans. In all iterations of cluster analysis, IVs remained low and average within-group homogeneity high within the higher-level cluster comprising Clusters 4-6. The clusters retained are those with the highest IVs within the higher-level cluster: Cluster 4 sites occur mainly on slopes (Table 1, p. 160). The widespread woodland species Xeroderris sthlmannii is the best indicator (Figure 2, p. 152). Pterocarpus erinaceus and Bombax costatum are also important structurally (Table 7, p. 167), but both have IVs<10% for this cluster. Cluster 5 is also dominated by generalists. No species has an IV>20%. Oxytenanthera abyssinica, Pterocarpus erinaceus, and Combretum glutinosum are the most important species (Table 8, p. 168). These sites are mainly in high topographic positions, with silty soil overlying a ferricrete hardpan (Table 1, p. 160). Most sites have undisturbed vegetation, but some were occupied or cultivated >30 y.a. Generalists also dominate Cluster 6 sites; only Combretum nigricans has an IV >20% (Figure 2, p. 152). These sites have a wide range of environmental characteristics: disturbed and undisturbed sites with nearly every substrate textural class belong to this cluster (Table 1, p. 160). Combretum glutinosum and Pterocarpus erinaceus are important structurally, as are the less widespread species Combretum nigricans and Hexalobus monopetalus (Table 9, p. 169), both of which are associated with poor, rocky soil (Arbonnier 2000).

124 Few sites belong to Cluster 7, which is strongly distinct from other clusters and has the lowest between-site heterogeneity of all clusters (Figure 2, p. 152) and relatively high diversity (Table 3, p. 163). These sites are all undisturbed, with non-arable silty to clayey soil (Table 1, p. 160). Gardenia ternifolia and Crossopteryx febrifuga strongly indicate Cluster 7 (Table 10, p. 170). Structurally, several widespread species that are uncommon in other clusters have high importance in Cluster 7. Terminalia macroptera strongly indicates and dominates Cluster 8 vegetation (Figure 2, p. 152; Table 11), but the tree is also common in several other clusters. Pterocarpus erinaceus and Combretum glutinosum are also important structurally. About half of Cluster 8 sites are undisturbed, and about half are abandoned fields (Table 1, p. 160). Clusters 9 and 10 both represent bushland vegetation dominated by Pterocarpus lucens and other Sahelian species. Different, strong associations with substrate texture distinguish these clusters: Exposed Sandstone B bedrock occupies 25% of the ground surface in all Cluster 9 sites (Table 1, p. 160). In these sites, patches of exposed bedrock are interspersed with grassy patches of shallow soil. Nearly all woody plants are rooted in bedrock fractures, not soil. The best indicator of this cluster is the cactiform succulent Euphorbia sudanica (Figure 2, p. 152). Many component species also occur in mesic, GilletiodendronHippocratea gallery forest (Cluster 2); their association with Sandstone B is observed also in Cluster 9 sites. Amongst these Sandstone B specialists is Combretum micranthum, which co-dominates Cluster 9 vegetation with Pterocarpus lucens (Table 12, p. 172). Gilletiodendron glandulosum is moderately important in Cluster 9 vegetation, especially in downslope portions of these sites.

125 Combretum micranthum and Pterocarpus lucens dominate in the drier, upslope conditions most characteristic of this cluster. Diversity is fairly high in this cluster (Table 3, p. 163) because it includes species with both upslope and downslope affinities. Cluster 10 includes all, and only, sample sites with clayey red (i.e. plinthitic) soil (Table 1, p. 160). These sites occur in naturally eroded areas near seasonal drainagechannel heads and have almost no herbaceous cover. Pterocarpus lucens and Guiera senegalensis strongly indicate this vegetation (Figure 2, p. 152), which is dominated structurally by Pterocarpus lucens and Combretum glutinosum (Table 13, p. 173). All Cluster 11 sites have disturbed vegetation, and all but one are abandoned settlements (Table 1, p. 161). Cluster 11 vegetation is physiognomically bushland, indicated and structurally dominated by Dichrostachys cinerea and Ziziphus mauritiana (Table 14, p. 174), both species characteristic of disturbed ground (Arbonnier 2000; Devineau 2005; Devineau 2001). Diversity is low in this cluster (Table 3, p. 163), but several economically important species are important or indicative, including the introduced, domestic tree Moringa oleifera (33.2%IV, p=0.003). Clusters 12-15 represent Sudanian woodland vegetation as classically conceived (e.g. Aubrville 1950; Chevalier 1938). These clusters are distinctive as a group, but are not strongly different from one another (Figure 2, p. 152). All four clusters are dominated by Pterocarpus erinaceus, Combretum glutinosum, and Terminalia macroptera, and useful trees are indicative or important (e.g. Vitellaria paradoxa, Adansonia digitata). Most sites are abandoned fields or settlements with deep, loamy soil without rock, and gentle slope (Table 1, p. 160). Importantly, no undisturbed sites were identified with similar soil and slope characteristics, but several undisturbed sites having shallow, rocky soil or strong slopes also belong to these clusters:

126 Vegetation in Cluster 12 sites is dominated by Pterocarpus erinaceus and Combretum glutinosum (Table 15, p. 175), but no species has an IV15% (Figure 2, p. 152). Compositional heterogeneity is moderate, and diversity low. Most sites are abandoned fields or settlements, and of these most are long-abandoned fields (Table 3, p. 163). However, eight undisturbed sites also belong to Cluster 12. Combretum micranthum, Acacia ataxacantha, and Hexalobus monopetalus are moderately important components of this vegetation. All three are characteristic of poor, rocky soil (Arbonnier 2000), suggesting that soil in these sites has low fertility.

The importance of Terminalia macroptera, Vitellaria paradoxa, and Piliostigma thonningii distinguish Cluster 13 vegetation (Table 16, p. 176), although Prosopis africana is the best indicator (Figure 2, p. 152). All Cluster 13 sites are disturbed, and nearly all have deep, sandy soil (Table 1, p. 160). Several economically important species (especially Vitellaria paradoxa) are important in Cluster 13 (and Cluster 15) vegetation.

Cluster 14 consists of both disturbed and undisturbed sites, with silty to sandy soil (Table 1, p. 160). Pteleopsis suberosa and Hymenocardia acidaboth small trees with wind-dispersed fruitare good indicators of this vegetation (Figure 2, p. 152), although the generalists Pterocarpus erinaceus and Combretum glutinosum are most important structurally (Table 17, p. 177). Most Cluster 14 sites are abandoned settlements, but many undisturbed sites and abandoned fields are also included.

Finally, Cluster 15 sites are dominated strongly by Pterocarpus erinaceus, which results in low diversity per site (Table 18, p. 178) and low between-site heterogeneity (Figure 2, p. 152). Combretum glutinosum and Vitellaria paradoxa are also

127 important structurally. Most Cluster 15 sites are abandoned settlements (Table 3, p. 163). This vegetation is similar to that of Cluster 13, except Terminalia macroptera is a minor component of Cluster 15 vegetation, and diversity is much lower in Cluster 15 (Table 3, p. 163). Ordination results. All three final solutions of NMS ordination had low instability but high stress relative to Kruskals (1964) and Clarkes (1993) guidelines for assessing NMS results. NMS stress is a measure of departure from monotonicity in the relationship between the dissimilarity (distance) in the original p-dimensional space and in the reduced kdimensional ordination space (McCune & Grace 2002: 125-126). The large size of these data sets is the probable source of the high stress: Kruskals and Clarkes guidelines are based on analysis of relatively small data sets (Kruskal & Wish 1978; McCune & Grace 2002). Stress increases for larger data sets, and if contributions to final stress are distributed roughly evenly across points, high-stress results can be interpreted with moderate reliability (Clarke 1993). In the present research, examination of the distribution of sites in ordination space and r-values for each species and all axes suggested that no site or species contributed an inordinately large portion of final stress. For the all-site NMS analysis, the most stable solution arranged sample sites along two axes (final stress=23.29, final instability=0.00051). In ordination space, sample sites and species occupy two swarms corresponding to the primary division in cluster analysis: forest versus woodland/bushland sites (Figure 3, p. 154). Clusters 1-3 segregate fairly clearly in ordination space, but few clusters in the woodland/bushland cluster are distinct. In the all-site ordination, the first axis explains a small portion of variation (r2=0.155), while the second axis explains much more (r2=0.403). (NMS ordination does not identify axes in order of strength of

128 correlation: McCune and Grace 2002.) Undisturbed-site ordination was the least stable of the three NMS analyses (final stress=21.29, final instability=0.0062). The first axis in this analysis explains a moderate amount of variation (r2=0.239), the second axis much more (r2=0.424) (Figure 4, p. 156). Forest clusters remain distinct in this ordination, but other vegetation clusters are less clearly defined in ordination space. Disturbed-site NMS ordination identified three axes in the most stable solution (final stress=21.30, final instability=0.0002). In this ordination, the first (r2=0.193), second (r2=0.245), and third (r2=0.232) axes each explain moderate amounts of variation (Figure 5, p. 158). Ruins and fallows weakly segregate in this ordination. Correlation coefficients for environmental factors and species distributions in relation to each of the seven ordination axes range between strongly negative to strongly positive (Figures 3-5, pp. 154-159; Table 19, p. 179; Appendix 1, p. 180). Based on observations of the microhabitats in which species were observed, as well as published information on species ecology, the ordination axes appear to relate to: All-site axis 1: Species with positive correlation coefficients are mesophytes, while those with negative correlation coefficients are xerophytes. All-site axis 2: Species with positive correlation coefficients are adapted to growth in rock outcrops, while those with negative correlation coefficients are most abundant in arable sites. Undisturbed-site axis 1: Species with positive correlation coefficients are mesophytes, while those with negative correlation coefficients are xerophytes. Undisturbed-site axis 2: Species with positive correlation coefficients are most abundant in arable sites, while those with negative correlation coefficients are adapted to growth in rock outcrops.

129 Disturbed-site axis 1: Species with positive correlations are most abundant in undisturbed sites with sandy loam to loamy sand, while those with negative correlation coefficients are most abundant in disturbed sites with silty loam. Disturbed-site axis 2: Species with positive correlations are most abundant in undisturbed sites with silty loam, while those with negative correlation coefficients are most abundant in disturbed sites with sandy loam to loamy sand. Disturbed-site axis 3: Species with positive correlations are most abundant in old fallows, while those with negative correlation coefficients are most abundant in new ruins. Discussion The results indicate three main causes of variation in vegetation composition in the research area: soil moisture, bedrock geology, and disturbance. Soil moisture. Topography and hydrogeology create habitat patches with permanently elevated soil moisture conditions. The two axes identified in the all-site and undisturbed-site ordinations relate to these two different sources of variation in soil moisture content. In both ordinations, the first axes relate to soil moisture determined by landscape position, as suggested by the r-values for hydrology (Figures 3-4, pp. 154-157). Also, the species with the highest positive correlations with these axes (Table 19, p. 179) are those indicative or characteristic of riparian bamboo forest (Cluster 3), which dominates low-lying sites along drainage channels with 25% of the ground surface covered by rocks >5 cm in diameter (Table 1, p. 160). Sites that lie toward the right end of this axis (as represented in Figures 3-4, pp. 154-157) have permanently moist soil conditions. The second axes in these ordinations relate to soil moisture conditions as determined by groundwater flow, as indicated by high r-values for the factor

130 substrate texture (Figure 4, p. 156). Sites with cobbly to bouldery soil are scree slopes below outcrops, where soil moisture is high because runoff is concentrated at the base of exposed bedrock (DCTD 1990). Sites with exposed bedrock have fractures where runoff collects or where perched aquifers are accessible (DCTD Dunne 1990; DCTD 1990). The species with the strongest, positive correlations to these axes are all characteristic or indicative of vegetation Clusters 1, 2, or 9 (Tables 4, 5, 12, & 19, pp. 164, 165, 172, & 179), which represent nearly all sites with extremely coarse substrate texture. High r-values for the factors slope and permanent water are indicative of spatial correlation for these factors: sites with exposed bedrock substrate most commonly occur on steeply sloped outcrops, and most permanent water sources are springs originating from between sedimentary layers in exposed sandstone bedrock (see Chapter 6). The high r-value for the factor disturbance is misleading, because disturbed sites swarm at one end of the second axes. In contrast, substrate texture explains the position of vegetation clusters along the entire length of these axes. Hydrogeography and topography both create permanently moist soil conditions, which is the primary cause of variation in vegetation composition across the landscape (cf. Breman & Kessler 1995; de Bie et al. 1998; Fournier 1991; Lawesson 1995), represented by the woodland/bushland versus forest split in the cluster analysis (Figure 2, p. 152). Bedrock and vegetation. Sudano-Guinean gallery forest (Clusters 1 and 2) occurs at seepage areas and on seasonal drainage channels along bedrock outcrops, where mesic conditions allow hygrophilous plants to survive. Since the 1930s, researchers have recognized these patchesespecially those dominated by Gilletiodendron glandulosum, endemic to the Manding Plateauas relict vegetation because many range-restricted or extralimital species occur in them (e.g. Aubrville 1939; Duong 1947; Jaeger 1956). However, few researchers have

131 noticed that many species characteristic of Gilletiodendron-Hippocratea forest (Table 5, p. 165) also occur in xeric, rupicolous bushland (Table 12, p. 172). These vegetation types are uniquely associated with exposed Sandstone B. Anthropogenic deforestation theory has dominated views of vegetation history in West Africa, and has until recently discouraged studies of edaphic causes of vegetation variation in the region (Cole 1986). The dominant view of vegetation history in Mali holds that GilletiodendronHippocratea forest patches are remnants of the presumed original forest climax that has been destroyed by allegedly poor African land management (Duvall 2003). To proponents of this view, forest patches occur only where topography protects vegetation from fire and human activities (e.g. Jaeger 1956; Jaeger 1968; Schnell 1976). Rupicolous scrub has not been specifically considered in the context of anthropogenic deforestation theory. While the inaccessibility of Gilletiodendron-Hippocratea forest sites contributes to their low level of direct human use (Duvall 2001; Geerling 1985), the present research shows that similarly inaccessible sites with different types of bedrock do not host Gilletiodendron-Hippocratea forest, but other types of Sudano-Guinean gallery forest. These results confirm a longstanding, alternative model of phytogeography in the Manding Plateau, which is poorly developed in West Africa but well supported by observations elsewhere. Proponents of this view propose that some vegetation types, especially gallery forest, are indicators of patchily distributed hydrogeological conditions. In 1917, the geologist Rn Chudeau proposed that fractures in sandstone bedrock capture and hold surficial runoff, which slowly seeps to the surface and allows azonal hygrophilous vegetation to survive. He made similar observations elsewhere in West Africa (e.g. Chudeau 1910; Chudeau 1913), and Larminat (1927) independently proposed this mechanism for plateaus in central Mauritania.

132 Subsequently, the botanist Paul Jaeger also concluded that sandstone hydrogeology is significant in the distribution of Gilletiodendron-Hippocratea forest, but he believed that forest vegetation creates mesic soil conditions, and not that groundwater availability allows forest vegetation to establish in mesic sites (e.g. Jaeger 1950b; Jaeger & Jarovoy 1952; Jaeger & Winkoun 1962). Ultimately, Jaeger embraced the anthropogenic theory of vegetation distribution (e.g. Jaeger 1956; Jaeger 1968; Jaeger 1966), which had been promoted by earlier botanists (e.g. Aubrville 1939; Duong 1947). Although the early geological observations have received no attention from botanists working in the Manding Plateau since Raynal and Raynal (1961), the biogeographic significance of perched aquifers in sandstone outcrops is well established from research conducted on other continents (Bowman et al. 1990; Bowman et al. 1988; Danin 1999; Davis 1951; Dunne 1990; Walck et al. 1996). The Manding Plateau is a geographic unit within the extensive West African sandstone massif that has distinct cultural and biogeographic features (Chudeau 1921; Jaeger 1959; Jaeger 1966), but it is not geologically distinct from other parts of the massif. Structural features of the sandstones comprising the massif result in predictably variable hydrogeology and erosion, which drive landform development (Chudeau 1917; Daveau 1959; Larminat 1927; Michel 1973; Urvoy 1942). The Manding Plateau comprises three sedimentary series overlying a weakly metamorphosed granitic basement complex (DNGM 1992; Groupement Manantali 1979; Varlet et al. 1977). Dolomite intrusions occur throughout the area. At a broad scale, perched aquifers in rock fractures distinguish the sedimentary rocks from the igneous rocks, which hold little groundwater internally (DCTD Chudeau 1917; DCTD 1990). However, the sedimentary series include various rocks with differing structural characteristics, which cause perched aquifers to be

133 distributed in predictable locations across the landscape. Simplistically, these sedimentary rocks can be divided into three broad groups. Most importantly, there are hard, well-cemented sandstonesincluding Sandstone Bthat form the distinctive cliffs of the Manding Plateau (Varlet et al. 1977). These sandstones have low intergranular porosity, but hold much groundwater in intercrossed bedding planes and strongly expressed vertical fractures (DCTD 1990; Groupement Manantali 1979). Second, in most areas, the hard sandstones overlie massive, weakly cemented sandstonesincluding Sandstone 0with indistinct, horizontal bedding planes and uniform grain size. Although these sandstones have high intergranular porosity, they lack well-developed networks of fractures that hold water, and serve mainly as barriers to vertical throughflow, especially when saturated (Groupement Manantali 1979). Finally, in some areas, thin layers of fine-grained rocks, mainly siltstone, interpose layers of the two broad types of sandstone described above. These fine-grained rocks have low porosity and permeability, and also serve as barriers to throughflow. In short, barriers to vertical throughflow cause the hard, well-cemented sandstones to serve as groundwater reservoirs. Groundwater flow at these barriers is horizontal; some water may penetrate the underlying rock where fractures occur, but such infiltration is generally low (Groupement Manantali 1979). Instead, the groundwater tends to scour tunnels along preexisting fractures in the hard sandstone, thus increasing the strength of horizontal flow (cf. Dunne 1990). Field observations suggest that oblique stratifications in Sandstone B direct groundwater flow to portions of the rock with horizontal stratifications (cf. Campbell 1973; Dunne 1980; Schick 1965). In several cases, cliff faces were observed where past fragmentation followed vertical fractures and the upper surface of strongly expressed stratifications, indicating a relative hardness that would impede throughflow. Indeed, in all sites in the research area where

134 groundwater permanently seeps from Sandstone B outcrops, stratifications are horizontal, and the strongest flow occurs at the uppermost stratum. Seepage erosion (sensu Dunne 1990) of the hard sandstones results in steep, angular outcrops due to disaggregation along vertical fractures and horizontal bedding planes. Many authors have described this geomorphic process throughout the West African massif (e.g. Daveau 1959; de Chtelat 1938; Jaeger & Jarovoy 1952; Larminat 1927; Michel 1973; Schnell 1960; Urvoy 1942). These authors have not observed that disaggregation is most active at the heads of gorges with groundwater seepage. Lithological features that concentrate groundwater flowsuch as fractures, or horizontal stratifications amid oblique stratalead to the extension of spring heads through seepage erosion, and to the formation of surface drainage networks (Campbell 1973; Dunne 1980). This distribution indicates that erosion by groundwater has created the deep, narrow gorges in which Gilletiodendron-Hippocratea forest often occurs (cf. Howard et al. 1988; Kochel et al. 1985). Rupicolous bushland occurs in adjacent, less steeply sloped areas of exposed bedrock upslope of a gorge head, where less runoff is available due to topography, but where plants adapted to Sandstone B can access groundwater via vertical fractures. Composition varies between Gilletiodendron-Hippocratea forest and rupicolous bushland because the former hosts mesophytes (e.g. Sarcocephalus latifolius, Spondias mombin) and the latter xerophytes (e.g. Pterocarpus lucens, Combretum glutinosum) that are not specially adapted to the physical structure of Sandstone B. Both Axes 1 of the all-site and undisturbed-site ordinations directly indicate correlation between species distribution and soil-moisture conditions, because these axes represent floristic variation determined by hydrology (i.e. landscape position). The second axes in these ordinations indirectly and only partly indicate species response to soil-moisture availability, but directly indicate specialized adaptation to the

135 most distinct plant habitat across the landscape, Sandstone B outcrops. For instance, many of the dominant species in Gilletiodendron-Hippocratea forest (Table 5, p. 165) have high, positive rvalues for the second axis of the undisturbed-site ordinations (Table 19, p. 179), even though these species occur in forest, woodland, and bushland clusters. In contrast, Pterocarpus lucens, indicative of rupicolous bushland, has a neutral r-value for the same axis (r=-0.006), but a relatively high r-value for the first axis (r=-0.41), indicating its drought tolerance. Pterocarpus lucens is not a sandstone specialist, but a species tolerant of dry soil conditions. The weakly cemented, porous sandstones of the Manding Plateau absorb water in interstices between grains. During the rainy season, these sandstones darken as they become saturated: the shade of one Sandstone 0 outcrop in the research area changed from 10 R 6/4 to 10 R 4/4 between 20 May and 25 September 2004. Additionally, fractures and junctures between strata, in which water may accumulate, are rare in these porous sandstones, as suggested by the characteristically rounded shape of Sandstone 0 outcrops. Although dolomite is harder and less porous, its hydrogeology is similar because it does not hold much groundwater in fractures (DCTD 1990). Thus, there are no sites of permanent groundwater flow associated with dolomite or Sandstone 0 in the research area. Seasonal seeps occur sparingly in Sandstone 0, but these have discharge rates sufficient to support only algae, mosses, and small ferns. SpondiasColaBombax gallery forest (Cluster 1) occurs on colluvial slopes below outcrops of all bedrock types. These sites also have elevated groundwater levels, but for different reasons from that described above for Gilletiodendron-Hippocratea forest. Runoff is channeled along the face of outcrops and collects underground along subsurface portions of the bedrock, whether the rock is permeable or not (DCTD 1990). Vegetation composition differs

136 between Clusters 1 and 2 because plants that are specialized to the physical structure of Sandstone B dominate Cluster 2 sites. Bedrock geology does not account completely for the distribution of the biogeographically notable species characteristic of Gilletiodendron-Hippocratea forest, including Gilletiodendron glandulosum. Several sites with Sandstone B bedrock that appear to be suitable for Gilletiodendron-Hippocratea forest instead host SpondiasColaBombax forest (Cluster 1). Fire and human activities certainly have roles in shaping the distribution of these forest types, as do historical and ecological factors such as dispersal, climate variation, and natural disturbance (Kellman & Meave 1997; Kellman & Miyanishi 1982; McCune & Allen 1985; Morison et al. 1948). However, past research that attributed the patchy distribution of Gilletiodendron-Hippocratea forest to anthropogenic disturbance has not shown that this vegetation type, or the species that comprise the vegetation, are singularly restricted to sites inaccessible to fire and humans, regardless of other characteristics. The present research shows that indicator species for Gilletiodendron-Hippocratea forest (Figure 2, p. 152) are restricted almost uniquely to Sandstone B outcrops (Tables 4-18, pp. 164-178). These species occur in easily accessible and fire-prone rupicolous bushland, and other types of vegetation occupy inaccessible, narrow gorges in dolomite outcrops and ferricrete crusts. Floristic patterns of human disturbance. All but one disturbed sites (110 of 111), as well as 70 of 106 undisturbed sites, belong to the woodland/bushland cluster (Figure 2, p. 152). Disturbed sites predominate in Clusters 11-15, but only Clusters 11, 13, and 15 are composed entirely of disturbed sites (Table 1, p. 160). The effects of settlement and cultivation on vegetation composition in the research area are difficult to characterize for three reasons. First, the factor human disturbance cannot be satisfactorily controlled in sampling design. No sites

137 were discovered that had deep, loamy soil, few rocks, and gentle slope, and no evidence of past cultivation or settlement. There are probably very few such sites in Sudanian West Africa as a whole (Schnell 1976; White 1983), and it is unjustifiable to present any Sudanian woodland vegetation as undisturbed without supporting historical evidence. Second, interpreting the disturbed-site ordination (Figure 5, p. 158), which suggests sources of variation in composition between disturbed sites, is difficult because the second two axes in this ordination do not strongly correspond to any measured environmental factor. Finally, there is insufficient information available on the ecology of many species to make strong conclusions about the meaning of correlations between species distributions and disturbed-site ordination axes (Table 19, p. 179). Within these constraints, the results suggest that vegetation response to anthropogenic disturbance is variable across the landscape. Disturbed sites as a group, or sub-groups based on type of or time since disturbance, are not homogenous (Table 2, p. 162). Most disturbed sites occupy the lower, central portion of allsite ordination space, which represents sites with sandy to loamy soil, gentle slope, and midslope topographical position (Figure 3, p. 154). However, disturbed sites are not closely grouped in ordination space. Similarly, disturbed sites are only broadly grouped in the dendrogram resulting from cluster analysis (Figure 2, p. 152). Human disturbance does not have a single, unambiguous effect on vegetation composition. The results show that the effects of cultivation and settlement on vegetation composition vary across the landscape, depending on biophysical factors and land-use practices. Cultural ecologists working in African woodlands have observed three responses to disturbance caused by cultivation or settlement. All three of these responses are evident in the research area:

138 Vegetation turnover occurs when disturbance increases the abundance of previously uncommon species, often increasing plant diversity through intentional or unintentional plant introduction. This outcome of disturbance has gained prominence through Fairhead and Leachs (1996) work in southern Guinea, in which they argue that farmers purposefully create forest patches at settlement sites by planting or protecting Guineo-Congolian rainforest species and thus increasing their abundance relative to Sudano-Guinean woodland species. Amanor (1994) described similarly teleological vegetation turnover in Ghana, while Avenard et al. (1974) and Spichiger and Pamard (1973) in Cte dIvoire and Duvall in Mali (see Chapter 5) report unintentional turnover (although the responsible farmers recognize and understand this process). Bush encroachmentin which woody shrub abundance increases relative to grasses in woodlands where early dry season fires become more frequent, often due to livestock management practicesis also appears a form of vegetation turnover (cf. Audru 1977; Bassett & Boutrais 2000; Bassett & Koli Bi 2000). In the research area, vegetation turnover occurs primarily at ruins, but also in some fallows. Fallows and ruins do not strongly segregate in cluster analysis (Table 1, p. 160; Figure 2, p. 152) or all-site ordination space (Figure 3, p. 154). This is because fallows and ruins generally occupy environmentally similar sitesthose with relatively deep, fertile soilwhere Sudanian woodland species dominate. However, fallows and ruins weakly segregate in the disturbed-site ordination (Figure 5, p. 158), because settlement and cultivation differ as ecological disturbances. Of course, settlement and cultivation varies between individuals and over time, and each settlement or field site has a distinct history. Thus, vegetation turnover does not

139 occur in all ruins, and may occur in some field sites. Dichrostachys-Ziziphus bushland (Cluster 11: Table 14, p. 174)the only cluster composed primarily of abandoned settlement sites (Table 1, p. 160)most clearly represents turned-over vegetation, although vegetation composition in Clusters 12-13 (Tables 15-16, pp. 175-176) also suggests turnover. Many species associated with vegetation turnover following settlement disturbance are strongly, negatively correlated to the three axes of the disturbed-site ordination (Table 19, p. 179), which also correlate strongly to past use (Figure 5, p. 158). Species associated with turnover have one or more of the following traits: a) edible fruits (e.g. Ximenia americana, Annona senegalensis), b) high economic value (e.g. Adansonia digitata, Moringa oleifera), or c) general adaptation to disturbance (e.g. Dichrostachys cinerea, Ziziphus mauritiana: Devineau 2001, 2005; Arbonnier 2000). These traits suggest that two processes, both related to plant dispersal, contribute to vegetation turnover. The main cause of turnover is the colonization of disturbed sites by postdisturbance specialists. Although vegetative reproduction is important in postdisturbance succession in fallows in semi-arid West Africa (Nyerges 1989), it is less important in ruins because settlement is a more intense disturbance than cultivation: a) few trees and no sucker sprouts are spared in settlement clearings, while many trees and sucker sprouts are maintained in fields; b) soil compaction is higher in settlements than fields; and c) settlements are occupied for more years and during more of each year than fields (see Chapter 2). This increases the importance of seed dispersal in vegetation succession following settlement abandonment. While many

140 widely dispersed, woodland species remain abundant in settlement sites following abandonment, vegetation in ruins is distinctive because species with high dispersal potentialthose adapted to colonizing woodland canopy openingsincrease in abundance. Colonizing species include anemochores (e.g. Pterocarpus erinaceus, Combretum glutinosum) and zoochores (e.g. Lannea velutina, Acacia seyal) that have relatively low economic value. Second, humans contribute directly to vegetation turnover by increasing the abundance of economically valuable species in settlement sites. While such species are often highly visible in and qualitatively indicative of ruins, individually they comprise only modest components in terms of vegetation structure. Vegetation in ruins includes introduced, domesticated trees: Moringa oleifera was the only such species observed in >5 sites, but mango (Mangifera indica) and cashew (Anacardium occidentale) were also encountered. More important, though, are economically valuable, African native trees, especially ronier palm (Borassus aethiopum) and baobab (Adansonia digitata). These valuable, native species are regionally widespread and do not appear to have narrow habitat preferences (Arbonnier 2000; Breman & Kessler 1995; Lawesson 1995), but in the research area are rare outside of settlements (see Chapter 5). The presence of these plants in ruins is due primarily to human activities; all have human-dispersed seeds and high use values, and are adapted to the edaphic conditions characteristic of settlement sites (see Chapter 5). Vegetation turnover is relatively easy to recognize, because it is indicated by the relative abundance of species that are uncommon elsewhere in the landscape. The

141 other two vegetation responseshomogenization and resilienceare more difficult to recognize. Vegetation homogenization occurs when species number decreases following disturbance, usually due to the elimination of rare species. This outcome of disturbance has been recognized for decades (e.g. Aubrville 1947; Chevalier 1928) and has been used to support the desertification and deforestation discourses that have dominated aspects of natural resource management in Africa (cf. Bassett & Crummey 2003; Leach & Mearns 1996). Nonetheless, Devineau (2005) has shown convincingly that homogenization occurs following cultivation in western Burkina Faso, similar to earlier findings in Sierra Leone (Nyerges 1989), Benin (Schreckenberg 1999), and Senegal (Lykke 1998). In the research area, homogenization occurs primarily in agroecologically marginal fallows, increasing the distinctness of fallows in the disturbed-site ordination (Figure 5, p. 158). Terminalia-Pterocarpus-Combretum woodland (Cluster 8: Table 11, p. 171) most clearly exemplifies homogenized vegetation although composition in Clusters 4-6 and 15 also suggests this process. Cluster 8 is composed of eight undisturbed sites and seven fallows (Table 1, p. 160), and has low plant diversity (Table 3, p. 163). None of the undisturbed sites are arable because of infertile or shallow soil, while the disturbed sites are arable. That vegetation composition in these sites is similar despite disturbance history suggests that human disturbance has reduced plant diversity in the fallow sites (Devineau 2005). While the fallow sites all have arable soil and gentle slope, they are considered poor for farming because their soil is fairly shallow, very sandy or silty, or relatively infertile

142 (see Chapter 3). The undisturbed sites have even lower agroecological potential: they are steeply sloped, and/or have rocky, silty, clayey, or shallow, infertile soil. Maninka farmers do not generally consider such sites farmable. Many species associated with vegetation homogenization following settlement are strongly, positively correlated to the three axes of the disturbed-site ordination (Table 19, p. 179), which also correlate strongly to past use (Figure 5, p. 158). These species are strongly associated with sandy loam and loamy sand (e.g. Terminalia macroptera, Albizia malacophylla), or silty loam (e.g. Crossopteryx febrifuga, Strychnos spinosa), further indicating that sites with poor, but arable, soil are most susceptible to homogenization following disturbance. Other species indicative of homogenized vegetation are associated with infertile soil, such as Combretum nigricans, Hexalobus monopetalus, and Acacia ataxacantha (Arbonnier 2000). Agricultural practices that deplete soil fertility to a greater degree than customary practiceincluding farming on agroecologically marginal sitesmay lead to vegetation homogenization (Nyerges 1989). Specific farming practices (e.g. length of cultivation, crops planted) were not assessed for each fallow site, and would be difficult to assess for many sites. In most cases, widespread generalists dominate homogenized vegetation (e.g. Terminalia macroptera, Pterocarpus erinaceus, Combretum glutinosum), contributing to low plant diversity. Homogenization results from reductions in the abundance of uncommon or specialist species (Devineau 2005). This does not mean that past human activities have homogenized all vegetation dominated by widespread generalists. Large areas of woodland vegetation with no history of settlement or cultivation occur in sites that are not arable. In the present research, undisturbed

143 woodland sites dominate the distinct, if heterogeneous, vegetation Clusters 4-7 (Table 1, p. 160). These sites (and others labeled undisturbed in this paper) are not pristine because they are exposed to low intensity and low frequency disturbance via hunting, livestock grazing, wild plant harvesting, and wild honey collection, but such activities do not move these sites a great distance toward the humanized pole of the disturbance continuum (cf. Vale 2002). The facile labels derived woodland and undifferentiated woodland, which have been applied to Sudanian woodlands to substitute for recognition of different woodland communities (Lawesson 1994; Lawesson 1995), mask variability in human impacts on woody vegetation (cf. Laris 2002). Vegetation resilience exists when disturbance has little effect on species number. Resilience has been reported in southern Africas Zambezian woodlands, where species composition remains stable over time despite repeated clearing (Chidumayo 2004; McGregor 1994; Stromgaard 1986). Resilience of woody vegetation to disturbance has not been reported from Sudanian woodland sites. The fact that settlement and cultivation have ambiguous and equivocal effects on vegetation composition suggests that Sudanian woodlands are resilient to disturbance. While seed dispersal by humans and other vectors alters the abundance of some species in some sites, the dominance of vegetative reproduction amongst woodland trees enhances ecosystem memory and allows similar communities to develop after disturbance as long as disturbance events do not exceed ecosystemspecific intensity and frequency thresholds (Nyerges 1989). In other words,

144 vegetation turnover and homogenization may be relatively short-lived vegetation responses to disturbance, at least in some cases. In the research area, resilience is clearest in the composition of PteleopsisPterocarpus woodland (Cluster 14: Table 17, p. 177). Several species associated with resilient vegetation have strong, positive correlation to the third axis of the disturbed-site ordination (Table 19, p. 179), which correlates to past use (Figure 5, p. 158). Such species include Pterocarpus erinaceus, Bombax costatum, and Anogeissus leiocarpus, which have long been considered indicative of climax vegetation in the Sudanian woodland zone (e.g. Aubrville 1949a; Aubrville 1949b; Chevalier 1938; Chevalier 1933; Schnell 1976; White 1965; White 1983). These species are abundant in both old and new fallows (and ruins) that have resilient vegetation. Vegetation resilience is also suggested more generally in the disturbed-site ordination. Age-classes of ruins and fallows are moderately clustered along the third axis of this ordination, and the oldest disturbed siteswhether fallows or ruinstend to occupy the central portion of ordination space (Figure 5, p. 158). The effects of settlement and cultivation on vegetation composition are ambiguous and equivocal, and the present results do not prove that vegetation turnover, homogenization, or resilience has occurred in any specific sample site or vegetation cluster. Additional research designed to assess specifically the effects of human disturbance on particular plant communities will be necessary to describe actual vegetation change in the research area. In contrast, the present research was meant to contextualize anthropogenic vegetation by situating it in the broader context of vegetation variation across the focal landscape. Furthermore, these

145 observations do not apply to herbaceous vegetation, which varies independently of woody vegetation (Devineau 2005). While much research remains in order to describe actual vegetation change in the research area, these results underscore that anthropogenic vegetation change in Sudanian West Africa is more complex than often represented. First, vegetation composition in ruins and fallows is not identical, because settlement and cultivation differ ecologically as disturbances. Few cultural ecologists working in West Africa have studied settlement as a distinct land use (see Chapter 2), and ecological understanding of rural settlements as woody plant habitat lags behind that of fields and fallows. Second, although specific vegetation clusters are cited above as indicative of each of the three vegetation responses, compositional data for most clusters that include many disturbed sites could be interpreted to suggest more than one response to disturbance. The direction and intensity of anthropogenic vegetation change varies between sites for social and ecological reasons (Vale 1982), and ecologically similar sites may express different vegetation responses if socioeconomic factors caused farming or settlement practices to differ, even subtly, between sites (Nyerges 1989). This aspect of disturbance was not assessed in the present research. Finally, observation of specific vegetation responses may be timedependant, if responses endure only for limited times following disturbance. The present results suggest that turnover and homogenization may be short-lived, although the length of time a site shows these responses probably varies between sites. Keeping in mind the complexity and variability of vegetation response to disturbance suggested here, the different responses to disturbance previously observed for African woodland sites may reflect different approaches to studying vegetation change as much as cultural ecological differences between research sites. Researchers who have observed similar vegetation

146 responses to disturbance in different sites have studied vegetation change in similarly limited portions of focal landscapes, and have focused on similar groups of species. Studies that have found vegetation resilience have focused on mean vegetation attributes in fallows (Chidumayo 2004; McGregor 1994; Stromgaard 1986). Mean vegetation attributes are determined largely by widespread, generalist species. In contrast, newer studies that show or suggest vegetation homogenization have focused on the abundance of uncommon, specialist species (e.g. Devineau 2005; Lykke 1998; McCune & Grace 2002; Nyerges 1989; Schreckenberg 1999). Finally, studies that have shown vegetation turnover have relied primarily on sampling in ruins or intensively managed fields, and not in less intensively disturbed fields and fallows (Amanor 1994; Fairhead & Leach 1996; Spichiger & Pamard 1973). Vegetation response across focal landscapes is likely more complex than suggested by studies of specific, limited areas. However, narrowly focused studies are necessary to reduce complexity and allow precise description of actual vegetation change. Conclusion Vegetation in southwestern Mali has interested botanists since the 1930s because the endemic tree Gilletiodendron glandulosum has been viewed as evidence for recent anthropogenic deforestation (Duvall 2003). However, this dominant representation of GilletiodendronHippocratea forest neglected Chudeaus (1917) earlier explanation that provided a possible biophysical explanation of Gilletiodendrons endemism and patchy distribution. The present research provides reason to resurrect the scientific discourse linking plant geography and bedrock hydrogeology that has been largely forgotten in the Africa context. This approach to understanding plant geography provides a fresh perspective for understanding how humans have affected vegetation composition in West Africa.

147 For instance, the case of Gilletiodendron-Hippocratea forest contrasts clearly with the better-known case of Polylepis forests in the Andes, another narrowly endemic habitat. The patchy distribution of Polylepis forest was considered natural, associated with specific types of terrain, until research in the 1980s showed that these patches are not restricted to a narrow range of topographic or edaphic situations (Purcell et al. 2004). Instead, biogeographic analysis has shown that Polylepis patches survive only in sites that are topographically protected from fire, grazing, and human activities (Fjelds 2002). In contrast, this research shows that the biogeographically notable species in Gilletiodendron-Hippocratea forest are restricted to a narrow range of sites with specific hydrogeological characteristics, and are not uniquely found in topographically protected sites. Gilletiodendron-Hippocratea forest and rupicolous bushland should be considered edaphic vegetation types, and not remnants of past deforestation. The dominant spatial structure of vegetation variation in the research area is attributable to edaphic variation, especially the hydrogeology of Sandstone B. Settlement and cultivation result in observable changes in vegetation characteristics, but only in those parts of the landscape with edaphic and hydrological conditions suitable for agriculture. The biogeography of Gilletiodendron-Hippocratea forest, and Sudanian West Africa more generally, must be understood in a global context. The bedrock-vegetation link described in the present paper helps create high-diversity patches of vegetation around the world, particularly mesic refugia in semi-arid areas (Bowman et al. 1990; Bowman et al. 1988; Danin 1999; Davis 1951; Walck et al. 1996; Woodford 2000). In short, the protective topography created by sharply angular, hard sandstone outcrops, and ultimately the resilient hydrogeology of these outcrops, create small patches of highly stable habitat that allows mesophytes to survive despite climate desiccation and human disturbance (Larson et al. 2000). Hard, sandstone outcrops must be more

148 generally recognized as important refuges for paleoendemics (Larson et al. 2000)exemplified by Wollemi pine (Wollemia nobilis W.G. Jones, K.D. Hill & J.M. Allen), a Tertiary relict discovered in Australia in 1994 (Woodford 2000). Recognition of the biogeographic importance of hard sandstone outcrops has significant implications for biodiversity conservation. Other parts of the West Africa sandstone massif also host a high number of biogeographically notable and endemic plants (e.g. Jaeger & Winkoun 1962; Porembski & Brown 1995; Schnell 1960). Management strategies that focus on protecting sites, rather than restricting activities, should be more effective to conserve these plants and the vegetation that hosts them (Danin 1999; Larson et al. 2000; Maxted et al. 1997). Better understanding of hydrogeology and biogeography in hard sandstone massifs is necessary for understanding how dispersed networks of refugial patchesnot just linear riparian forest patches (cf. Meave & Kellman 1994; Meave et al. 1991)buffer the impacts of climate change on biodiversity.

149 Figures, tables, and appendix for Chapter Three

150 Figure 1. Maps of West Africa, western Mali, and the research area. Map 1 shows West Africa including the West African sandstone massif (after Jaeger & Winkoun, 1962), and the location of the area shown in map 2. Map 2 shows western Mali, including the location of the research area. Map 3 shows the research area, including the location of Solo.


152 Figure 2. Dendrogram produced by cluster analysis. Similarity scale given at top. Physiognomic categories shown at the left of dendrogram follow Lawessons (1995:24) definitions. Average between-site distance given as x. For indicator values (IVs), p < 0.001 unless another value is given.


154 Figure 3. All-site NMS ordination. Cluster numbers (in ordination plot at right) refer to vegetation clusters represented in Figure 2 (p. 152) and described in Tables 4-18 (pp. 164-178).


156 Figure 4. Undisturbed-site ordination. Abbreviations: bushl. = bushland; woodl. = woodland. For species abbreviations, see Appendix 1 (p. 180).


158 Figure 5. Disturbed-site NMS ordination.


160 Table 1. Environmental factors. Abbreviations: n=sites per environmental factor; undist.=undisturbed (no evidence of past clearing); woodl.=woodland; bushl.=bushland; cult.=cultivated; ya=years ago; sett.=settled. Plite (French) means fine-grained sedimentary rock. In some analyses, factors were simplified by lumping categories. Thus, permanent water lumped the first four landscape position categories and maintained the fifth. The following factors were measured quantitatively: past use (by time since abandonment, lumping the first three categories, and others according to time, not use), substrate texture (by estimating mean grain size for soil or colluvial rock, and considering bedrock the greatest grain size), and slope (by degrees). Landscape position was quantified by estimating average distance per site to drainage channel or permanent spring (after IGN 2001 and field data); bedrock was quantified by ranking rocks by increasing hardness (i.e. plite, ferricrete, Sandstone 0, Sandstone B, dolomite).


Environmental factor Forest, undist. Woodl., undist. Bushl., undist. Cult. 10 ya Cult. 10-20 ya Cult. 20-30 ya Cult. 30 ya Sett. 10 ya Sett. 10-20 ya Sett. 20-30 ya Sett. 30-40 ya Sett. 40 ya Clayey loam Silty loam Loam Sandy loam, sand 25% cobbles 25% boulders 25% bedrock Topo. high Topo. low Any plot 50m from channel All plots 100m from channel Any plot 100m from spring dolomite ferricrete plite Sandstone 0 Sandstone B 2 5 10 >10 Past use Subsrate texture Hydrology Bedrock Slope

n 34 52 20 17 14 14 16 5 7 12 11 15 18 34 30 87 11 15 22 97 54 41 16 9

1 13

2 11

3 10 2 1

4 7 1

5 9

6 6 4

7 4

8 8

9 6

10 10


12 8


14 8


1 3 1 1 2 6 2 2

2 2 2 1

1 4 2 5 4 7 2 4 2 2 5 2 11 1 3 3 4 1 1 11 1 10 1 10 8 3 2 1 2 6 3 6 2 1 10 2 1 2 4 8 1

1 2 6 1 2 1 3 10 2 2

1 3

10 2 5 5 2 1 11 4 6 6 10

3 1 3 1 1 2 4 2

2 1 2 4 2 1 5 4 7 3 3 4 24 1 1 11 16 9

3 2 1 2 2 2 1 1 1 12 1 2 7 5

3 3 5 1 1 3 1 1 6 5 15

4 6 4 7 1 1 2 6 19


2 2

4 2 2 1

16 9 1

8 12 5

10 79 2 2 1 28 98 10 132 31 9 3 45 10

9 3 8 1 3

1 6 7 8 3 3

6 3 6 8 2 5 6 3 2 1


1 5 3 7 2

3 1

7 3

2 10 1 8 15 34 1 1

1 1 4 8 12 2

1 15 1 2 7 19 3 4

10 8 7 22 3

162 Table 2. MRPP results. Statistics: T=test statistic, more negative values indicating greater between-group heterogeneity; A=agreement statistic, within-group homogeneity increases as A approaches 1; p=statistical significance.
Grouping basis Cluster results Binary: disturbed/undisturbed Binary: presence/absence of permanent water Sites grouped by disturbance type, time since disturbance, and substrate texture; undisturbed sites further grouped by vegetation structure Substrate texture (as shown in Table 1, p. 160) Landscape position (as shown in Table 1, p. 160) Slope categories (as shown in Table 1, p. 160) Disturbed sites grouped by time since disturbance; undisturbed sites lumped Disturbed sites grouped by time since and type of disturbance; undisturbed sites lumped Geological parent material (as shown in Table 1, p. 160) T -69.07 -61.99 -53.06 -46.43 -44.48 -39.64 -31.48 -30.30 -25.56 -10.98 A 0.71 0.16 0.14 0.42 0.29 0.21 0.15 0.18 0.21 0.07 P <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001

163 Table 3. Vegetation cluster characteristics. Abbreviations: med.= median; dist.=between-site distance (Srenson [Bray-Curtis] index). Cluster numbers correspond to those shown in Figure 2 (p. 152). Berger-Parker diversity index equals the proportional abundance of the most abundant species per site.
Cluster Number of individuals All Per sites site (SD) 1237 95.2 (37.4) 1357 121.4 (27.6) 1096 84.3 (39.5) 339 33.1 (16.7) 679 56.6 (17.3) 457 32.7 (13.3) 210 49.5 (9.0) 1188 81.6 (41.5) 208 32.5 (14.1) 653 66.3 (15.6) 796 81.1 (33.4) 2031 56.4 (18.1) 1071 76.5 (29.5) 2474 91.9 (33.7) 2843 114.2 (35.7) Number of species All Per sites site (SD) 80 19.5 (6.2) 53 17.2 (5.0) 64 13.9 (5.7) 43 11.5 (4.3) 49 17.5 (2.9) 63 12.9 (3.5) 22 12.0 (1.8) 45 12.9 (4.4) 29 11.7 (3.9) 37 14.6 (2.1) 53 15.8 (3.7) 80 13.7 (3.7) 53 15.9 (3.2) 75 19.5 (5.0) 64 16.0 (3.9) Number of families All Per sites site (SD) 34 14.8 (3.6) 31 13.6 (3.1) 29 11.2 (3.7) 21 7.5 (2.5) 22 12.0 (1.8) 31 8.9 (2.7) 11 7.0 (1.4) 22 8.9 (3.1) 20 9.2 (3.1) 16 8.8 (1.6) 27 11.1 (2.5) 33 9.6 (2.5) 25 10.9 (2.4) 33 12.2 (2.8) 32 11.2 (2.5) Berger-Parker diversity index Range Med. 0.120.52 0.170.35 0.300.82 0.120.46 0.140.47 0.180.39 0.160.26 0.230.66 0.170.43 0.270.55 0.210.56 0.190.55 0.130.40 0.130.38 0.190.76 0.19 0.23 0.59 0.31 0.27 0.25 0.21 0.4 0.22 0.33 0.37 0.30 0.22 0.23 0.44 Mean dist. Total sites Total area (m2) 13,300 11,000 12,000 9,000 12,000 14,000 4,000 15,000 6,000 10,000 9,000 36,000 14,000 26,000 25,000

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15

0.27 0.04 0.15 0.22 0.18 0.41 0.03 0.07 0.14 0.08 0.14 0.14 0.15 0.10 0.08

13 11 12 9 12 14 4 15 6 10 9 36 14 26 25

164 Table 4. Dominant species for Cluster 1 (Cola cordifoliaSpondias mombinBombax costatum forest). For species abbreviations, see Appendix 1 (p. 180). Proportional abundance=individuals per species divided by total individuals. Relative frequency=sites per species divided by total sites. Basal dominance=basal area (m2) per hectare. All species shown having proportional abundance 0.05 or relative frequency 0.50. Top five species shown for basal dominance.
Proportional abundance Boco 0.17 Spmo 0.09 Dime 0.07 Oxab 0.07 Grbi 0.06 Coco 0.05 Relative frequency Spmo 1.00 Dime 0.92 Coco 0.92 Comi 0.77 Boco 0.77 Oxab 0.69 Grbi 0.62 Sala 0.62 Stku 0.62 Feap 0.62 Algl 0.54 Maal 0.54 Basal dominance Spmo 2.08 Oxab 2.04 Coco 1.37 Stku 1.20 Comi 0.99

165 Table 5. Dominant species for Cluster 2 (Gilletiodendron glandulosumHippocratea indica forest). For explanation of table contents, see Table 4, p. 164.
Proportional abundance Gigl 0.23 Hiin 0.19 Boco 0.09 Grbi 0.08 Sase 0.07 Comi 0.05 Relative frequency Gigl 1.00 Hiin 1.00 Grbi 1.00 Comi 0.91 Spmo 0.91 Boco 0.82 Sase 0.82 Sala 0.73 Oxab 0.73 Coto 0.73 Boan 0.73 Diab 0.64 Stsa 0.64 Gyam 0.55 Dime 0.55 Basal dominance Gigl 11.55 Boco 2.26 Spmo 2.25 Oxab 1.76 Addi 1.48

166 Table 6. Dominant species for Cluster 3 (Oxytenanthera abyssinica [Bamboo] forest). For explanation of table contents, see Table 4, p. 164.
Proportional abundance Oxab 0.54 Xest 0.05 Rasu 0.05 Relative frequency Oxab 1.00 Sala 0.75 Pter 0.67 Fisu 0.58 Khse 0.50 Rasu 0.50 Anle 0.50 Sase 0.50 Basal dominance Oxab 22.82 Ersu 0.52 Khse 0.31 Rasu 0.27 Cepe 0.18

167 Table 7. Dominant species for Cluster 4 (Xeroderris sthlmanniiPterocarpus erinaceus Bombax costatum wooded grassland). For explanation of table contents, see Table 4, p. 164.
Proportional abundance Xest 0.19 Boco 0.15 Pter 0.13 Oxab 0.05 Relative frequency Pter 1.00 Xest 0.89 Boco 0.89 Cogl 0.89 Tema 0.78 Oxab 0.56 Hemo 0.56 Basal dominance Addi 2.15 Pter 1.83 Oxab 1.32 Boco 1.14 Xest 0.76

168 Table 8. Dominant species for Cluster 5 (Ferricrete wooded grassland). For explanation of table contents, see Table 4, p. 164.
Proportional abundance Oxab 0.16 Cogl 0.14 Pter 0.07 Acat 0.06 Relative frequency Pter 1.00 Cogl 0.92 Laac 0.83 Oxab 0.75 Boco 0.67 Demi 0.67 Tema 0.67 Hemo 0.67 Lave 0.58 Acat 0.58 Stsp 0.58 Vima 0.58 Buaf 0.50 Xest 0.50 Doqu 0.50 Selo 0.50 Basal dominance Oxab 15.98 Pter 1.31 Boco 1.03 Cogl 0.68 Lave 0.68

169 Table 9. Dominant species for Cluster 6 (Combretum nigricansHexalobus monopetalus wooded grassland). For explanation of table contents, see Table 4, p. 164.
Proportional abundance Hemo 0.12 Coni 0.09 Cogl 0.08 Pter 0.06 Relative frequency Cogl 0.93 Hemo 0.71 Pter 0.64 Coni 0.50 Tema 0.50 Basal dominance Addi 2.60 Pter 1.20 Anle 0.96 Hemo 0.38 Laac 0.31

170 Table 10. Dominant species for Cluster 7 (Crossopteryx febrifugaGardenia ternifolia wooded grassland). For explanation of table contents, see Table 4, p. 164.
Proportional abundance Tela 0.19 Crfe 0.12 Gate 0.12 Cogl 0.10 Demi 0.09 Ptsu 0.07 Pter 0.06 Daol 0.05 Relative frequency Buaf 1.00 Crfe 1.00 Demi 1.00 Enaf 1.00 Gate 1.00 Laac 1.00 Tela 1.00 Cogl 0.75 Daol 0.75 Ptsu 0.75 Basal dominance Pter 3.13 Demi 2.74 Laac 2.70 Tela 2.47 Buaf 1.44

171 Table 11. Dominant species for Cluster 8 (Terminalia macropteraPterocarpus erinaceus woodland). For explanation of table contents, see Table 4, p. 164.
Proportional abundance Tema 0.46 Pter 0.14 Cogl 0.05 Relative frequency Tema 1.00 Pter 1.00 Cogl 0.87 Vipa 0.87 Xest 0.73 Laac 0.60 Stsp 0.53 Boco 0.53 Basal dominance Oxab 4.64 Tema 3.20 Pter 2.79 Laac 1.38 Copi 0.62

172 Table 12. Dominant species for Cluster 9 (Rupicolous bushland). For explanation of table contents, see Table 4, p. 164.
Proportional abundance Comi 0.15 Hemo 0.10 Eusu 0.09 Cogl 0.09 Gigl 0.09 Pter 0.06 Grbi 0.06 Ptlu 0.06 Hiin 0.05 Relative frequency Ptlu 1.00 Cogl 0.83 Pter 0.83 Boan 0.67 Comi 0.67 Eusu 0.67 Grbi 0.67 Hemo 0.67 Boco 0.50 Gigl 0.50 Gyam 0.50 Hiin 0.50 Lami 0.50 Basal dominance Ptlu 0.74 Cogl 0.42 Gyam 0.35 Grbi 0.24 Comi 0.16

173 Table 13. Dominant species for Cluster 10 (Pterocarpus lucensGuiera senegalensis bushland). For explanation of table contents, see Table 4, p. 164.
Proportional abundance Ptlu 0.29 Cogl 0.16 Comi 0.11 Acat 0.06 Hemo 0.05 Relative frequency Cogl 1.00 Ptlu 1.00 Acat 0.80 Hemo 0.80 Laac 0.80 Boan 0.60 Casi 0.60 Guse 0.60 Pter 0.60 Tela 0.60 Comi 0.50 Coni 0.50 Copi 0.50 Daol 0.50 Stsa 0.50 Xiam 0.50 Basal dominance Ptlu 1.45 Oxab 0.75 Cogl 0.65 Afaf 0.42 Xiam 0.35

174 Table 14. Dominant species for Cluster 11 (Dichrostachys cinereaZiziphus mauritiana bushland). For explanation of table contents, see Table 4, p. 164.
Proportional abundance Dici 0.33 Zima 0.13 Cogl 0.10 Pter 0.06 Pith 0.05 Alma 0.05 Relative frequency Dici 1.00 Zima 0.89 Cogl 0.78 Pith 0.78 Pter 0.78 Acse 0.56 Addi 0.56 Alma 0.56 Boae 0.56 Doqu 0.56 Hemo 0.56 Tema 0.56 Vipa 0.56 Basal dominance Addi 4.88 Oxab 1.44 Boae 1.16 Zima 0.76 Fisy 0.58

175 Table 15. Dominant species for Cluster 12 (Acacia ataxacanthaCombretum micranthum Hexalobus monopetalus woodland). For explanation of table contents, see Table 4, p. 164.
Proportional abundance Pter 0.30 Cogl 0.11 Zima 0.06 Pith 0.05 Dici 0.05 Tema 0.05 Relative frequency Pter 1.00 Cogl 0.94 Tema 0.69 Vipa 0.67 Pith 0.56 Zima 0.50 Basal dominance Addi 7.99 Pter 3.83 Oxab 3.29 Anle 1.21 Pith 0.56

176 Table 16. Dominant species for Cluster 13 (Terminalia macropteraVitellaria paradoxa Piliostigma thonningii woodland). For explanation of table contents, see Table 4, p. 164.
Proportional abundance Tema 0.15 Pter 0.13 Cogl 0.10 Vipa 0.07 Pith 0.06 Dici 0.06 Alma 0.06 Tela 0.05 Praf 0.05 Relative frequency Tema 1.00 Pith 1.00 Cogl 0.93 Pter 0.93 Vipa 0.93 Lave 0.86 Praf 0.64 Alma 0.57 Anse 0.57 Boco 0.57 Dici 0.57 Ptsu 0.50 Selo 0.50 Zima 0.50 Basal dominance Addi 5.35 Oxab 3.00 Vipa 1.78 Pter 1.19 Khse 0.99

177 Table 17. Dominant species for Cluster 14 (Pteleopsis suberosaHymenocardia acida woodland). For explanation of table contents, see Table 4, p. 164.
Proportional abundance Pter 0.18 Ptsu 0.13 Cogl 0.11 Tema 0.09 Hyac 0.05 Relative frequency Cogl 1.00 Ptsu 1.00 Pter 1.00 Lave 0.85 Tema 0.85 Hyac 0.81 Vipa 0.81 Crfe 0.65 Demi 0.62 Anse 0.58 Boco 0.58 Buaf 0.58 Pith 0.54 Stsp 0.54 Xest 0.54 Laac 0.50 Tela 0.50 Basal dominance Pter 5.73 Oxab 2.55 Tema 1.54 Addi 1.48 Vipa 1.44

178 Table 18. Dominant species for Cluster 15 (Pterocarpus erinaceusVitellaria paradoxa Piliostigma thonningii woodland). For explanation of table contents, see Table 4, p. 164.
Proportional abundance Pter 0.36 Cogl 0.23 Pith 0.06 Vipa 0.05 Relative frequency Cogl 1.00 Pter 1.00 Vipa 0.84 Lave 0.76 Tema 0.76 Pith 0.64 Zima 0.56 Ptsu 0.52 Alma 0.48 Hyac 0.48 Basal dominance Pter 10.91 Oxab 3.56 Vipa 2.34 Addi 1.88 Cogl 1.82

179 Table 19. Correlations between species distributions and ordination axes. The highest and lowest five Pearsons r correlation coefficients for each axis of the three NMS analyses are shown. For Pearsons r correlation coefficients for all species in all ordinations, see Appendix 1 (p. 180).
All-site ordination Axis 1 Axis 2 Ptlu -0.39 Pter -0.42 Hemo -0.38 Cogl -0.37 Crfe -0.37 Vipa -0.36 Laac -0.35 Lave -0.33 Demi -0.31 Zima -0.31 Ptsa 0.28 Sala 0.51 Fisu 0.28 Hiin 0.51 Sala 0.33 Gigl 0.54 Khse 0.34 Grbi 0.63 Oxab 0.35 Spmo 0.65 Undisturbed-site ordination Axis 1 Axis 2 Cogl -0.58 Grbi -0.69 Ptlu -0.41 Gigl -0.63 Laac -0.40 Hiin -0.61 Hemo -0.38 Spmo -0.58 Tela -0.37 Gyam -0.50 Ersu 0.43 Vipa 0.36 Coto 0.46 Crfe 0.39 Sala 0.54 Buaf 0.41 Oxab 0.55 Tema 0.46 Spmo 0.57 Pter 0.53 Disturbed-site ordination Axis 1 Axis 2 Axis 3 Laac -0.56 Zima -0.58 Zima -0.51 Addi -0.40 Dici -0.41 Pith -0.50 Hemo -0.36 Addi -0.38 Dici -0.39 Xiam -0.34 Fisy -0.33 Anse -0.37 Mool -0.32 Acse -0.31 Mool -0.32 Cole 0.29 Crfe 0.44 Boco 0.24 Alma 0.36 Demi 0.47 Pter 0.34 Praf 0.40 Stsp 0.47 Stsp 0.37 Pith 0.46 Hyac 0.50 Oxab 0.38 Tema 0.54 Ptsu 0.56 Anle 0.47

180 Appendix 1. Species abbreviations and correlation coefficients for ordination axes. Abbreviations: spp.=species; Ax1, etc.=Axis 1, etc.; undist.=undisturbed (no evidence of past settlement or cultivation). No correlation coefficients are provided for species that occurred in <5 disturbed or undisturbed sites. Nomenclature follows Hutchinson and Dalziel (1954-1972) or Geerling (1982).

All-site ordination Ax1 Ax2 r r -0.29 -0.07 0.17 -0.20 0.16 -0.21 -0.17 -0.03 0.11 0.32 0.25 0.30 0.15 -0.23 0.13 0.15 0.17 -0.07 -0.10 -0.28 0.02 0.06 0.14 -0.20 -0.17 0.44 0.11 0.41 0.15 -0.04 0.04 0.00 -0.30 -0.15 -0.25 -0.01 0.20 0.31 0.18 0.28 0.25 0.45 -0.12 -0.37 0.14 -0.15 -0.18 0.36 -0.11 -0.06 0.22 0.15 -0.21 0.14 -0.24 -0.17 0.27 0.50 -0.37 -0.20 -0.21 -0.13 -0.31 -0.18 0.12 0.39 0.18 -0.25 0.23 0.42 -0.04 -0.13 -0.20 -0.16 0.28 0.37 -0.16 0.17 0.03 0.26 0.14 0.38 -0.03 0.26 0.08 -0.11 0.05 0.14 0.28 0.26 0.27 0.01 -0.30 -0.16 Undist.-site Disturbed-site ordination ordination Ax1 Ax2 Ax1 Ax2 Ax3 r r r r r -0.34 0.17 -0.25 0.16 0.09 0.07 -0.31 -0.20 -0.40 -0.38 -0.22 0.24 -0.24 0.36 -0.18 0.36 -0.10 -0.18 0.11 0.15 0.07 -0.15 0.47 -0.15 0.29 0.15 0.30 -0.37 0.09 -0.04 -0.18 0.00 0.10 -0.27 -0.28 -0.27 -0.02 -0.43 0.33 -0.37 0.23 0.32 0.24

Spp. codes Acat Acse Addi Afaf Alco Algl Alma Anaf Anle Anse Bamu Boae Boan Boco Brfe Brmi Buaf Casi Cepe Cipo Coco Cogl Cole Comi Como Comy Coni Copi Coto Crfe Daol Demi Diab Dici Dime Doqu Enaf Ersu Eusu Feap Fiab Fico Fidi Figl Fisu Fisy Gate

-0.30 -0.20

0.41 -0.03 0.07 -0.02

0.42 0.16

0.23 0.20

0.31 -0.33 0.42 -0.33 -0.58 0.30 -0.11 -0.43

0.16 -0.09 -0.06 0.07 0.29 0.09 -0.01 0.01 -0.21

0.29 -0.11 -0.20 0.01 0.10 0.08 0.20 -0.34 0.31 -0.22 0.14 0.00 0.46 -0.38 -0.34 0.39 -0.10 0.44 0.01 -0.26 0.16 -0.25 0.36 -0.10 -0.32 0.34 0.05 0.47 -0.03 0.27 -0.36 -0.12 -0.41 -0.39 0.42 -0.37 -0.07 0.20 -0.14 0.06 -0.01 -0.33 0.28 -0.16 0.31 -0.08 0.43 -0.18 -0.16 -0.15 0.17 -0.24 0.31 -0.33

0.09 -0.14 0.38 -0.09 -0.32

0.04 -0.04 -0.26

0.07 -0.33 -0.15 0.30 -0.19 0.13 -0.13

Full species name Acacia ataxacantha DC. Acacia seyal Del. Adansonia digitata L. Afzelia africana Smith ex Pers. Allophyllus cobbe (L.) Raeusch Albizzia glaberrima (Schum. ex Thonn.) Benth. Albizzia malacophylla (A.Rich.) Walp. Antiaris africana Engl. Anogeissus leiocarpus (DC.) Guill. & Perr. Annona senegalensis Pers. Baissea multiflora A.DC. Borassus aethiopum Mart. Boscia angustifolia A.Rich. Bombax costatum Pellegr. & Vuillet Bridelia ferruginea Benth. Bridelia micrantha (Hochst.) Baill. Burkea africana Hook. f. Cassia sieberana DC. Ceiba pentandra (L.) Gaertn. Cissus populnea Guill. & Perr. Cola cordifolia (Cav.) R.Br. Combretum glutinosum Perr. ex DC. Combretum lecardii Engl. & Diels Combretum micranthum G.Don Combretum molle R.Br. ex G.Don Cordia myxa L. Combretum nigricans Lepr. ex Guill. & Perr. Cordyla pinnata (Lepr. ex A.Rich.) Milne-Redhead Combretum tomentosum G.Don Crossopteryx febrifuga (Afzel. ex G.Don) Benth. Daniellia oliveri (Rolfe) Hutch. & Dalz. Detarium microcarpum Guill. & Perr. Diospyros abyssinica (Hiern) F.White Dichrostachys cinerea (L.) Wight & Arn. Diospyros mespiliformis Hochst ex A.DC. Dombeya quinqueseta (Del.) Exell. Entada africana Guill. & Perr. Erythrophleum suaveolens (Guill. & Perr.) Brenan Euphorbia sudanica A.Chev. Feretia apodanthera Del. Ficus abutifolia (Miq.) Miq. Ficus cordata Thunb. Ficus dicranostyla Mildbr. Ficus glumosa Del. Ficus sur Forssk. Ficus sycomorus L. Gardenia ternifolia Schum.

Gigl Grbi Grfl Grla Guse Gyam Hemo Hiin Hyac Isto Khse Laac Lami Lave Lola Maal Mase Mool Oxab Pabi Pacu Pela Pith Praf Psse Pter Ptlu Ptsa Ptsu Rasu Sala Sase Scbi Selo Spmo Stku Stsa Stse Stsp Tasp Tela Tema Vido Vima Vipa Xest Xiam Zima Zimu 0.06 0.06 0.00 -0.05 -0.24 0.02 -0.38 0.07 -0.19 0.22 0.34 -0.35 0.05 -0.10 0.16 0.18 -0.04 0.13 0.35 0.07 -0.01 -0.22 0.12 0.07 0.03 0.03 -0.39 0.28 -0.20 0.26 0.33 0.16 0.01 -0.15 0.28 0.26 -0.07 -0.18 -0.25 -0.09 -0.18 -0.12 0.27 -0.19 0.04 -0.01 -0.23 0.24 0.19 0.54 0.63 -0.11 -0.02 0.01 0.43 0.04 0.51 -0.18 0.26 0.17 -0.14 0.26 -0.33 -0.02 0.37 -0.22 -0.17 0.27 -0.11 -0.10 -0.09 -0.23 -0.23 -0.11 -0.42 0.08 0.22 -0.25 0.07 0.51 0.35 -0.05 -0.21 0.65 0.32 0.34 -0.02 -0.15 -0.12 -0.18 -0.21 0.16 -0.14 -0.36 0.04 -0.18 -0.31 0.05 0.25 0.30 -0.07 0.02 -0.28 0.15 -0.38 0.24 -0.21 0.32 0.37 -0.40 0.14 -0.14 0.11 0.30 -0.63 -0.69 0.17 -0.24 -0.01 0.09 0.02 0.00 -0.04 -0.50 0.07 -0.36 0.27 -0.61 0.20 0.10 0.50 -0.14 0.00 0.27 -0.56 0.33 -0.20 0.33 0.07 0.37 0.06 0.11 -0.22 -0.29 -0.01 0.17 -0.32 -0.21 0.09 -0.30 0.00 0.02 -0.03 0.18 0.03 0.23 0.23 0.46 -0.09 0.13 0.40 0.17 0.53 0.15 0.03 -0.06 -0.05 0.31 -0.01 0.56 0.09 -0.38 -0.37 0.01 -0.14 0.08 -0.01 0.20 0.22 0.38 -0.58 -0.21 0.20 0.13 -0.29 0.09 0.30 0.03 0.47 0.00 -0.06 0.23 0.26 0.34 0.46 0.54 0.16 Gilletiodendron glandulosum (Port.) J.Lonard Grewia bicolor Juss. Grewia flavescens Juss. Grewia lasiodiscus K. Schum. Guiera senegalensis J.F.Gmelin Gyrocarpus americanus Jacq. Hexalobus monopetalus (A.Rich.) Engl. & Diels Hippocratea indica Willd. Hymenocardia acida Tul. Isoberlinia tomentosa Craib & Stapf Khaya senegalensis (Desr.) A.Juss. Lannea acida A.Rich. Lannea microcarpa Engl. & K.Krause Lannea velutina A.Rich. Lonchocarpus laxiflorus Guill. & Perr. Malacantha alnifolia (Bak.) Pierre Maytenus senegalensis (Lam.) Exell. Moringa oleifera Lam. Oxytenanthera abyssinica (A.Rich.) Munro Parkia biglobosa (Jacq.) Benth. Parinari curatellifolia Planch. ex Benth. Pericopsis laxiflora Harms. Piliostigma thonningii (Schum.) Milne-Redhead Prosopis africana (Guill. & Perr.) Taub. Psorospermum senegalense Engl. & Diels Pterocarpus erinaceus Poir. Pterocarpus lucens Lepr. ex Guill. & Perr. Pterocarpus santalinoides L'Hr. ex DC. Pteleopsis suberosa Engl. & Diels Raphia sudanica A.Chev. Sarcocephalus latifolius (Smith) Bruce Saba senegalensis (A.DC.) Pichon Sclerocarya birrea (A.Rich.) Hochst. Securidaca longipedunculata Fresen. Spondias mombin L. Stereospermum kunthianum Cham. Strophanthus sarmentosus DC. Sterculia setigera Del. Strychnos spinosa Lam. Tapinanthus sp. Terminalia laxiflora Engl. Terminalia macroptera Guill. & Perr. Vitex doniana Sweet Vitex madiensis Oliv. Vitellaria paradoxa Gaertn. f. Xeroderris sthlmannii (Taub.) Mendona & E.P.Sousa Ximenia americana L. Ziziphus mauritiana Lam. Ziziphus mucronata Willd.

-0.16 0.20

-0.08 0.00

-0.03 -0.06 0.15 -0.20 -0.32 0.38 -0.16 -0.22 -0.50 -0.14 0.34


-0.15 0.04 -0.12 -0.23 -0.41 0.35 -0.24 0.23 0.54 0.28 -0.25 0.57 0.39 0.02 -0.12 -0.22 -0.37 -0.18 -0.20 -0.04 0.11 -0.31 0.14


-0.15 0.06 -0.10 -0.01

0.37 -0.01 -0.16 0.20

0.30 0.04 0.39 0.22 0.36 0.09 0.18 -0.08 0.22 0.04 0.30 0.01 0.18 -0.34 0.19 -0.12 -0.16 -0.58 -0.51 0.01

183 Chapter Five: Human settlement and baobab distribution in southwestern Mali

Abstract Researchers have long assumed that human settlement establishment and reproduction of the baobab tree (Adansonia digitata) are spatially and temporally dependent because baobabs are abundant in many settlement sites in semi-arid Africa. This paper tests the spatiotemporal relationship between baobab and settlement distribution. In a study area of 183 km2, 1240 baobabs were located and mapped, their diameters measured, and habitat characteristics recorded for each individual. Second, all occupied (n=7) and abandoned (n=80) settlements were located and mapped, and occupation dates for each site were determined through interviews. Ethnographic observations revealed local knowledge of human-baobab interaction. Chi-square tests indicated baobab habitat preferences, and bivariate point-pattern analysis tested independence of observed point patterns of baobabs and settlements, including paired point sets consisting only of certain settlement age-classes and baobab sizeclasses. Statistical analyses support local knowledge of baobab-settlement attraction. Specifically, baobabs and settlements are attracted at most distances and for most baobab sizeclasssettlement age-class pairs. This attraction is significant only at distances of less than c.500 m. Young settlements are not significantly associated with large baobabs. Attraction between small and large baobabs is marginally significant at distances of less than c.500 m, but observed significance is less than that observed for attraction between baobabs and settlements. There are five main conclusions: 1) Human settlement and baobab recruitment are spatially dependant. 2) Settlement leads directly and indirectly to the development of baobab

184 groves at settlement sites. 3) Human activities cannot account for baobab presence in many parts of the landscape. 4) Although the baobab occurs in a wide range of habitats, its recruitment and mortality are not evenly distributed across these habitats. Recruitment is strongest in settlements, fields, and cliffs or steep slopes along rock outcrops, while mortality is highest on cliffs or steep slopes. 5) These habitat preferences suggest that baobab abundance in settlements is not caused simply by human seed dispersal, but also by other aspects of settlement practice that ensure dry, fire-protected settlement sites.

Introduction Long-term conservation and management of biodiversity resources depend on our ability to situate human activities within ecosystem processes (Micheli et al. 2001). Due to the urgency of addressing short-term threats to biodiversity resources, research on anthropogenic environmental change has focused on human activities having obvious effects at relatively short timescales, such as years to decades. However, in semi-arid Africa, humans have actively manipulated plant populations through management practices for millennia (O'Brien & Peters 1998; van der Veen 1999). Through much of the 20th century, recognition of this long history of resource management by African farmers and herders was expressed in terms that assigned blame for supposed environmental destruction on the supposed destructiveness of indigenous practices (Bassett & Crummey 2003; Fairhead & Leach 1998; Leach & Mearns 1996; Richards 1985). Dominant narratives of widespread, anthropogenic deforestation in West Africa have proven inaccurate or unsupportable (Bassett & Koli Bi 2000; Duvall 2003; Fairhead & Leach 1996; Ribot 1999). Nonetheless, land management practices do alter vegetation (Maranz & Wiesman 2003; Nyerges 1989; Nyerges 1997; Schreckenberg 1999; Turner 1998a; Turner

185 1998b). The emerging challenge is to develop land-use ecologies that account for observed variation in vegetation characteristics without resorting to oversimplified or inappropriate explanatory frameworks (Turner 2000). Humans are significant factors in the regeneration of many wild plant populations through activities that affect seed dispersal and germination (Maranz & Wiesman 2003; O'Brien & Peters 1998). In situations where markets exert strong influence on fruit use, humans may overharvest fruits and remove seeds from ecosystems, negatively impacting reproduction of target species and plant-frugivore interactions (Moegenburg 2002; Peres et al. 2003; Pruetz 2002). Where market influence is weaker, humans disperse seeds within ecosystems by harvesting and consuming wild fruits, which serve as important dietary items for people and domestic animals. In such situations, wild fruit seeds accumulate at settlement sites (Reid & Ellis 1995; van der Veen 1999). Fruit processingincluding selection, cleaning, cooking, and digestiondestroys some seeds, but also may promote germination in seeds that are not destroyed (Esenowo 1991; Johansson 1999; von Maydell 1992). Additionally, people often select and manage self-sown wild fruit trees in occupied and abandoned settlements and fields (Assogbadjo et al. 2005; Lovett & Haq 2000; Maranz & Wiesman 2003). As a result of fruit use and fruit tree management, wild fruit trees often dominate vegetation in fields, fallows, and settlements. In areas where these practices are combined with relatively intensive land use, in which most arable land is actively managed and fallow periods are fairly short, savanna parkland vegetation, dominated by economically important trees, develops (Boffa 1999; Etkin 2002; Neumann et al. 1998; Pullan 1974; Raison 1988). Although regionally widespread and characteristic of much of northern sub-Saharan Africa, parkland vegetation does not occur in many landscapes where land use has been

186 primarily extensive (Boffa 1999; Pullan 1974). Extensive farming has long been, and remains, important throughout semi-arid Africa (Raynaut 1997). In landscapes with long histories of land-extensive farming, the cumulative effects of human activities on indigenous plant distributions are unclear. There are well-worn assumptions about the destructiveness of landextensive agriculture, but these are often poorly founded on empirical assessment of agroecological dynamics (Dove 1983; Richards 1985). Humans have altered plant distributions at global, continental, regional, and local scales, but at local scalesacross landscapes measured in tens of kilometersthere is poor understanding of the biogeographic and ecological processes that lead to human-plant associations (Binggeli 1996; Cronk & Fuller 2001). The present paper uses point pattern analysis of tree and settlement distribution to assess the role of shifting settlement on the distribution of the baobab, a highly valued fruit tree, in southwestern Mali. Focal species The African baobab (Adansonia digitata L.) occurs in most of tropical Africas woodland areas, and its high utility as a source of food and various raw materials causes it to be one of the most salient trees for rural Africans (Dhillion & Gustad 2004; Kristensen & Lykke 2003; Owen 1970). Similarly salient to botanists due to its characteristic growth habit, the tree has attracted scientific attention for centuries; literature on the baobab is extensive (reviews include: Simpson 1995; reviews include: Wickens 1982). Nonetheless, knowledge of the trees ecology is remarkably thin (Assogbadjo et al. 2006; Assogbadjo et al. 2005; Baum 1995; Bowman 1997; Ebert 2006; Simpson 1995; Wickens 1982). Considering the trees large size, including large flowers and fruit, and gregariousness in many areas, the baobabs significance in African ecosystems is certainly underrepresented in present ecological knowledge.

187 The baobab appears to be an important link between the social and ecological processes that together create the effects of human activities on African ecosystems. Baobab has been part of agroforestry systems in West Africa for at least 1000 years (Kahlheber 1999; Neumann et al. 1998). Agricultural management practices and uses of baobab fruit, leaves, and seedlings affect the viability of baobab sub-populations (Assogbadjo et al. 2005; Baumer 1994; Dhillion & Gustad 2004; Johansson 1999). Indeed, baobabs dominate vegetation in and around settlements in many parts of semi-arid Africa (Boffa 1999; Enjalbert 1956; Rosevear 1937; Seignobos 1980; Wickens 1982). Indeed, abandoned settlements are often identifiable as baobab groves (Hobley 1922; Perron 1926; Sikes 1972; Taylor 1960). Apparent attraction between baobabs and settlements could arise in three ways. First, humans appear to be an important dispersal vector for the tree (Assogbadjo et al. 2006; Chevalier 1906; Dhillion & Gustad 2004; Guy 1971; Hobley 1922; Ridley 1930), so that the abundance of baobabs at settlement sites may result from fruit use. Second, humans may choose to establish settlements at preexisting baobabs for improved access to the resources these trees represent (Wickens 1982). Place names suggest that this relationship may be widespread. For instance, dozens of settlements named Sitakta Manding-language locative noun meaning under the baobab treeexist from Senegal to Burkina Faso (Office of Geography 1965a; 1965b; 1965c; 1966; 1965d). Third, settlement-baobab attraction may arise simply because human settlements and baobabs thrive in edaphically and topographically similar locations: the tree is most common on deep, well-drained soils that are also good for farming (Barnes 1980; Johansson 1999; Simpson 1995; Wilson 1988). In any case, researchers have not shown that human-baobab interactions are, indeed, spatially dependent.

188 Studies of baobab dispersal and reproduction have focused on individuals propagated or protected by humans in humanized vegetation, particularly settlement, field, and fallow sites (Dhillion & Gustad 2004; Johansson 1999), or have not explicitly considered land-use characteristics (Assogbadjo et al. 2006; Assogbadjo et al. 2005; Barnes 1980; Wilson 1988). Human activities are important in baobab reproduction, but it is necessary to contextualize human-baobab interactions by studying the trees ecology throughout focal landscapes. Scant information exists on its distribution or abundance in non-humanized vegetation, although several authors have identified several non-human dispersal agents, including large primates like chimpanzees (Pan troglodytes) and baboons (Papio spp.) (Ridley 1930; Wickens 1982). The baobab has no clear habitat or soil preferences, although it is generally most abundant on deep, sandy, well drained soils in arid to semi-arid woodlands (Simpson 1995; Wilson 1988). The trees aboriginal range may have been only drier parts of sub-Saharan Africa; its presence in more humid areas may result ultimately from human introduction, although it reproduces without human intervention and occurs in apparently undisturbed vegetation in semi-arid to sub-humid areas (Wickens 1982). There is no information on how the trees autecology, its interaction with non-human pollination and dispersal agents, and human activities together create spatial patterns of abundance in specific landscapes. Research area Research was conducted in an area of 183 km2 around Solo village in Malis Bafing Biosphere Reserve (Figure 1, p. 212). The research area approximately represents the area where residents of Solo, and no other villages, have traditional usufruct (although this customary tenure has no legal recognition). Solo has been occupied for about 400 years (Samak et al.

189 1986), and Solos current residents retain oral history of settlement for the last c.200 years (see Chapter 2). Biophysical setting. The research area lies in Malis Manding Plateau, where numerous sandstone outcrops rise 100-200 m above surrounding lowlands. The research area is bisected by the edge of a large sandstone plateau, with similar vegetation occurring above and below the plateau edge (Figure 1, p. 212). Vegetation is characteristic of Whites (1983) Sudanian regional center of endemism, dominated by woodland but with dispersed patches of forest, scrub, and grassland. The distribution of most vegetation types is linked to specific edaphic conditions: forest types occur in sites with moist soil conditions; woodlands and bushlands occur on relatively dry and deep, arable to non-arable soil; and grasslands occupy shallow soil and seasonally flooded areas (see Chapter 4 and Breman & Kessler 1995; Lawesson 1995). Human activities have ambiguous and equivocal effects on vegetation composition, but more clearly affect the distribution of some species (see Chapter 4). The abundance of useful trees, including baobab, is positively correlated with disturbance caused by settlement and cultivation. Elephants, which cause elevated baobab mortality in parts of East Africa (Barnes 1980; Barnes 1985; Caughley 1976; Weyerhaeuser 1985), were extirpated in the research area in about 1984, and were never abundant in the living memory of local residents (Duvall & Niagat 1997). All parts of the landscape are subject to at least low intensity or low frequency disturbance although evidence of anthropogenic disturbance is not readily apparent in many areas (Duvall 2001). The indigenous Maninka people settle and cultivate lowland sites with arable soil and good drainage. Rocky areas, steep slopes, sites with poor or shallow soil, and seasonally inundated areas are used only for seasonal livestock grazing, wild plant and honey collection, and hunting (Duvall 2001; Samak et al. 1987). Around settlements, farmers

190 annually burn most grassland and woodland areas to prevent destructive fires and to prepare fields (Laris 2002). Wild fruits and other plant products provide important food resources (Samak et al. 1987). During field clearing and subsequent management, farmers preserve individuals of several tree species with edible fruits, including baobab (Koenig & Diarra 1998; Samak et al. 1987). Farming is seasonal and rain-fed. Although farming practices vary between farmers and over time, individual fields are generally cultivated <10 years before fallowing >10 years (Samak et al. 1987). Arable soil is patchily distributed (PIRT 1983), and farmland is limited immediately around Solo. Thus, many farmers improve their access to farmland by establishing farming hamlets some distance from Solo, but within Solos area of traditional usufruct (see Chapter 2). Most hamlets are occupied only during the farming season and only for relatively short time periodsin most cases, <20-30 yearsbefore abandonment (Samak et al. 1986). Hamlets are usually occupied by only a small number of related, nuclear families who return to Solo after a hamlet is abandoned, and often establish other hamlets after some time in Solo. The practices of hamlet establishment and abandonment represent a shifting settlement system (cf. Stone 1996). Hamlet farming has probably been practiced in the research area for at least several hundred years, but has become increasingly important over the last century. Conservationists working in Mali consider hamlet farming a spatially uniform and destructive threat to natural habitat in the area (e.g. Caspary et al. 1998; PREMA 1996), but the cultural ecology of hamlet farming has not been studied. Methods

191 Data collection. Field research was conducted January-December, 2004, and entailed: 1) an ethnographic study of baobab-human interaction; 2) a census of baobab individuals, and 3) a census of settlement sites. Ethnographic study. Ethnographic research was conducted during January-December 2004 to understand the cultural ecology of settlement around Solo. Participant observation provided experiential knowledge of human-baobab interactions, while informal interviews clarified observations (Werner & Schoepfle 1987). Informal interviews were conducted by the researcher in the Maninka language, while working with farmers in fields, fallows, and settlements. Ethnographic data was analyzed in a qualitative manner (cf. Cotton 1996). Baobab census. The baobab census was based on interviews of local residents, and foot surveys. Only baobabs >1 m high were tallied. Interviews of c.45 peoplemales and females, c.10-80 years oldindicated the approximate locations of c.1000 baobabs. With the assistance of interviewees and other residents, the researcher located these baobabs. Additionally, local residents assisted the researcher in locating all abandoned settlement sites, which were searched for baobabs by walking a spiral path from the center of the site to a distance from the center of about 200 m. In occupied settlements, baobabs were located in collaboration with residents. Every occupied and abandoned settlement was visited and searched at least twice. Finally, foot surveys were conducted in areas: a) similar to those where many baobabs had already been observed; and b) where no baobabs or abandoned settlements were known. These foot surveys extensively covered the research area, including all drainage channels, edges of bedrock outcrops, edges of ferricrete hardpans, and known, active or abandoned paths. These efforts did not locate every baobab in the research area; the smallest DBH classes are certainly underrepresented. Nonetheless, a conservative estimate is that 95% of individuals were

192 located. For statistical analyses, the baobabs identified in the research area are considered a mapped, rather than sampled, point pattern (Bailey & Gatrell 1995). The location of each baobab was recorded using a Garmin GPS 12XL unit, and every tree was observed directly by the researcher. For each tree, the following data were collected: diameter at breast height (DBH), and habitat. DBH was measured using a diameter tape 5m long. For individuals with circumference at breast height >5m, circumference was measured using a 50-m tape, and diameter was determined arithmetically. Each individual >1m DBH (except those hosting beehives) was measured twice, during a single visit; the mean of the two measurements was recorded. The DBH of inaccessible individuals (e.g. on cliff ledges) was visually estimated. Habitat was simplified as comprising three environmental factors (Table 1, p. 226): a) topography, recorded either as slope angle (measured with an inclinometer), or as drainage channel, for individuals occurring along seasonal or permanent waterways; b) land use (i.e. occupied settlement or field, abandoned settlement or field, or no history of past use), determined from observable features or interview data (see Chapter 4); and c) vegetation, determined by structure (i.e. forest, woodland, scrub, thicket), with occupied fields and settlements lumped together as a single class. Additional vegetation characteristics (e.g. bamboo thicket, cliff-side gallery forest) provide specific information about vegetation ecology based on a concurrent vegetation sampling effort (see Chapter 4). No age estimates for baobabs were made because: a) DBH measurements were taken throughout the year, and baobab DBH varies significantly with short-term rainfall variation (Fenner 1980); b) age estimates from baobab DBH measurements do not appear to be robust in most cases (Johansson 1999); and c) local informants knew the ages of very few trees independently of the estimated ages of associated settlements or fields. However, DBH is used

193 as a general proxy for age because small individuals are considered younger than larger individuals (cf. Bowman 1997). Settlement census and history. A census of occupied and abandoned settlements was based mainly on interviews. Abandoned settlements were identified during three group interviews of Solos traditional authorities (chief, land chief, and senior counselors), all men aged c.40-80 years. Aerial photographs from 1952 and historical documents did not identify any additional settlement sites. With the assistance of interviewees, the researcher visited all occupied and abandoned settlement sites, sampled vegetation in sites that had not been subsequently farmed or occupied (see Chapter 4), and searched each site for baobabs. This data set is also considered a mapped point pattern (Bailey & Gatrell 1995). The location of each site was determined using a Garmin GPS-12XL unit, and was recorded as a point corresponding to the approximate center of the occupied area of each site (as evidenced by the distribution of huts or remains of hut foundations). For every site, the following information was collected: site name, estimated dates of establishment and abandonment, and subsequent use (i.e. cultivation, occupation, or subsequently undisturbed by cultivation or occupation). Generally, establishment and abandonment dates were estimated by correlating informant life history markers, changes in site occupation status, and datable events, such as national elections. Multiple informants were interviewed to triangulate date estimates and increase precision (cf. Flowerdew & Martin 1997). In some cases, specific dates of past site occupation were gathered from published documents (Anonymous 1958; de Lannoy de Bissy 1882; Park 1954 [1815]; Projet Inventaire 1990) or aerial photos from 1952. Since the goal of the present analysis is to determine if baobabs in a

194 settlement site precede or follow human occupation, only establishment dates were used in analyses. Although age estimates were established for the establishment and abandonment of each settlement, these age estimates were not used precisely in analyses because comparable estimates were not possible for observed baobabs. Instead, as with baobabs, the settlement data set was divided into quintiles based on estimated date of settlement establishment (Table 2, p. 227). Data analysis. Two analyses were used. First, baobab habitat preferences were assessed using chi-square tests (cf. Bowman 1997). Based on the size of each population quintile (248 individuals) and the number of environmental factor categories (Table 1, p. 226), the expected and observed numbers of baobabs per quintile per factor were compared (Bailey 1995). Second, the spatial independence of distribution patterns for baobabs and settlements was tested using Ripleys bivariate K function for several subsets of the data. There are numerous technical descriptions of the bivariate K function, which is calculated from the observed number of points (such as baobab individuals) in a given distance, h (e.g. Bailey & Gatrell 1995; Diggle 2003; Dixon 2002; Haase 1995). In application, the K function is generally linearized to stabilize variance and facilitate interpretation, and in this form is called the L function (Dixon 2002). Since the purpose of the present application is to examine the interaction of two point processes, significance testing was based on random toroidal shifts of the observed point patterns (Dixon 2002). Similar use of the bivariate K function to test spatial independence of two distribution patterns include Camarero et al. (2005), Eccles et al. (1999), Plissier (1998), and Couteron and Kokou (1997). Spatial analyses were conducted using the SPLANCS package (version 2.01) in the R statistical software environment (version 2.2.1). For further description of SPLANCS, see

195 Rowlingson and Diggle (1993), Gatrell et al. (1996), Bivand and Gebhardt (2000), and Diggle (2003). Multiple pairs of baobab and settlement point sets were used in bivariate L-function analyses (Table 2, p. 227). For each pair of baobab and settlement point sets, the bivariate L function was calculated using the equation L(h)=((K(h))/) h (Diggle 2003), with edge effects corrected geometrically (Bailey & Gatrell 1995). Values of L(h)>0 indicate attraction between types of points, while L(h)<0 indicates repulsion (Dixon 2002). There is no method for assessing significance for empirical L(h) values from non-rectangular study areas using toroidal shifts, and the present study area is not rectangular. Thus, significance of empirical LBaobabSettlement(h) values for the study area as a whole was not determined. Instead, three rectangular areas within the study area were designated in a post hoc manner to allow significance testing (Figure 2, p. 214). Bivariate analyses were conducted for each pair of baobab and settlement point sets in these rectangles; empirical LBS(h) values were compared with an envelope derived from 99 random toroidal shifts of data points (cf. Bailey & Gatrell 1995; Diggle 2003; Dixon 2002). Results Settlement practice and baobabs. Maninka farmers recognize that settlement leads to increased abundance of wild fruit trees, including baobab, at settlement sites. These increases are considered to result from interactions between tree species and humans, livestock, and wild animals. Maninka farmers clearly recognize the consequences of human activities on baobab abundance, but do not consider increased baobab abundance a planned outcome of settlement practice. Human-baobab interaction centers on the high material value of baobab products. Baobab fruit and leaves are probably the most frequently used wild foods. Women use baobab

196 fruit for flavoring breakfast porridge by dissolving the dry, sour pulp in hot water, then straining out the seeds and fiber embedded in the pulp. Briefly soaking seeds in boiling water stimulates germination (Esenowo 1991). Baobab seeds, discarded in rubbish mounds, are extremely abundant in the soil of occupied settlements and may remain dormant for many years (Baum 1995; Hobley 1922; Palmer & Pitman 1972), as illustrated in Figure 3 (p. 216). During fruiting season (November-January), successful women collect and store hundreds of fruits. Women also seasonally collect large quantities of young baobab leaves, which are powdered and used in preparing sauces for grain dishes, and men collect, harvest, and process baobab bark to make rope throughout the year. Due to the importance of these uses, men and women individually know the precise location of dozens of baobab trees, and often manage highly valued trees away from settlements or fields by clearing grass to reduce a trees vulnerability to fire damage. The baobabs non-material meanings increase its sociocultural significance. First, baobabs maintain history by physically representing past settlement (cf. Peluso 1996). Historical primacy is an important basis for socially granted resource-use rights (cf. Shipton 1994), which help determine an individuals access to natural resources (Ribot & Peluso 2003). Baobab trees may be named for the individuals who originally claimed rights to harvest their produce, usually by protecting self-sown trees. Baobab groves are recognized as, and called by the name of, settlements that indicate a familial groups historical primacy in a given area. Second, baobabs carry spiritual meanings. Baobabs are associated with beneficent spirits who may help advance a humans desires, and thus are places where people may safely invoke spiritual assistance without exposing ones wishes to malevolent spirits. Nonetheless, humans must interact cautiously with baobabs to avoid offending associated, benevolent spirits and risking their retribution.

197 These material, historical, and spiritual meanings all contribute to a strong prohibition against killing baobabs. When clearing vegetation for fields or settlements, baobab is one of three specieswith Vitellaria paradoxa C.F. Gaertn., and Parkia biglobosa (Jacq.) R.Br. ex G.Donthat informants describe as never being felled. However, new settlements are rarely established under preexisting baobabs, because harvesting rights for most large trees in arable sites have been determined previously. Maninka tree tenurewhich has no legal recognition is based upon a trees value and its origin in relation to human activities. Individual usufruct for trees of low-value or highly abundant species are generally not recognized, while individual usufruct for high-value trees are recognized if a person can reasonably claim to have been crucial to a trees establishment. Thus, men and women protect self-sown baobab saplings in fields or settlements in order to gain heritable harvesting rights. Nonetheless, since baobab trees fruit abundantly only after several decades (>30 years, according to Baumer 1983)that is, after its human owner, and often his/her children, are able to harvest the fruitmany baobabs are openly accessible resources even though their historic harvesting rights are respected in many situations, especially settlement and field establishment. Men establishing settlements avoid conflicts with preexisting rights to ensure that they will have unquestioned rights to any trees that sprout in their new settlement. Maninka settlements are habitat patches where wild fruit seeds accumulate and may find suitable conditions for germination. Settlement sites are selected based on several biophysical criteria that are determined through examination of topography, soil, and vegetation characteristics. The most significant criteria in terms of tree ecology are: a) deep, well drained soil, ideally sandy loam; b) good surface drainage; and c) the water table is shallow enough to access via hand-dug wells (i.e. <15-20 m). Sites with shallow or fine-grained soil, or poor

198 drainage are rejected for settlement because they are unsuitable for most crops, and because it is difficult to maintain mud buildings on moist soil. In establishing a settlement, male occupants clear all vegetation, except for about one moderately sized tree per family, to provide shade, and any Vitellaria, Parkia, or baobab individuals. Sites with very large trees are generally avoided, because they are difficult to fell, can attract pest birds, and pose a potential hazard during windstorms. During site occupation, people, aided by livestock, continuously clear herbaceous vegetation (to reduce snake and rodent habitat), and cut volunteers of low-value tree species (to reduce bird habitat). Volunteers of highly valued trees are spared, but generally not watered or protected with fencing. Browsing livestock, and leaf and bark harvesting severely injure and stunt most baobabs near settlements (cf. Arbonnier 2000). Although innumerable baobab seeds germinate in rubbish heaps each rainy season few survive; successful saplings are those that happen to grow in protected microenvironments (Figs. 3b, 3c). Informants recognize that baobab abundance increases after settlement abandonment because trees are no longer subject to humanand livestock-caused injuries. Other trees, particularly Vitellaria and Borassus aethiopum Mart., show increases in abundance similar to baobab, according to informants. Following site abandonment, baobabs grow quickly to become primarily sources of fruit rather than leaves or bark. Their growth, along with the growth of other trees surviving site occupation, initiates interactions between plants and animals that Maninka men, in their gendered role as hunters, recognize as transforming abandoned settlements into sites with an abundance of wild fruit trees (Figure 3d, p. 216). First, large trees shade the soil and create relatively cool and moist microenvironments (cf. Belsky et al. 1993). Second, chimpanzees and baboons frequently visit fruiting baobabs during the early dry season (November-January). While foraging, these animals trample the grass under baobabs they visit, which inhibits fire

199 from burning immediately under these trees. As a result, tree seedlings under productive baobabs are often spared from fires throughout the dry season. Large baobabs serve as nurse trees (cf. Simpson 1995), especially for plants with seeds dispersed by wild frugivores including Ficus spp., Tamarindus indica L., Spondias mombin L., and baobab. Ethnographic evidence further suggests that chimpanzees and baboons are important dispersal agents for baobab seeds. Hunters believe that most large baobabs at abandoned settlement sites are the result of human activities. However, they believe that nearly all small baobabs outside of occupied settlements and fields sprout from chimpanzee or baboon scat. This belief is based on the widely shared observations of Maninka hunters that: a) chimpanzees and baboons more frequently consume baobab fruit than any other wild animal; b) baobab seeds occur in chimpanzee and baboon feces; and c) small baobabs are abundant in habitats, especially along cliffs, where chimpanzees and baboons are most frequently seen, and where humans rarely visit. Baobab population structure. The baobab census identified 1240 individuals, a density of 6.69/km2 in the research area (Figure 2, p. 214). This density is similar to those reported elsewhere in Sudanian Africa: 5.0/km2 for northern Benin (Assogbadjo et al. 2005), 10.7/km2 for central Mali, and 11.2/km2 for southern Sudan (Wilson 1988). Other published density estimates range upward to 72.8/km2 (Weyerhaeuser 1985), with the highest densities occurring in northern Tanzania. The largest individual observed had a DBH of 455.0 cm, while 52 individuals were observed with DBH 1 cm. Notably, the average DBH of observed trees declined with the number of trees observed (Figure 4, p. 218), even though observations began in the dry season, when baobab DBH shrinks due to water loss, and continued through the rainy season, when

200 baobab DBH increases due to water gain (Fenner 1980). A histogram of DBH distribution (Figure 5, p. 219) suggests a Type III survivorship curve (Deevey 1947), which is characteristic of r-selected species, including most trees (Barbour et al. 1999; Pianka 1970). Similar size-class distributions have been reported for sampled baobab populations in Tanzania (Barnes 1980; Barnes 1985; Weyerhaeuser 1985). Elsewhere, baobab populations have exhibited bell-shaped size-class distributions (Assogbadjo et al. 2005; Caughley 1976; Johansson 1999; Wilson 1988). Habitat characteristics. Baobabs were observed in sites having a range of slope, vegetation, and land-use characteristics. However, baobabs are not evenly distributed with respect to the observed range of habitats; the tree is most abundant in occupied and abandoned settlements and fields. Additionally, some size classesespecially the largest and smallest individualsare significantly overabundant or underabundant in certain habitats (Table 1, p. 226). Recruitment and mortality are unevenly distributed. In particular, small baobabs are significantly overabundant in occupied settlements, occupied fields, and cliff faces (Figs. 3, 6), and underabundant on moderate slopes and in woodland vegetation. Large baobabs are significantly overabundant in old fields, and underabundant in occupied settlements and on cliff faces. Baobab spatial ecology. Habitat preferences revealed through chi-square analysis account for the patchiness of baobab distribution. Several large clusters occur throughout the research area, particularly at settlement sites (Figure 2, p. 214). These dense, broadly ovate clusters contrast clearly with the smaller number of generally linear and sparsely occupied clusters located along the edges of rock outcrops. Different size classes have different distributions across the landscape, but all size classes display a similar patchiness (Figure 2, p. 214). There are also several large gaps in baobab distribution that correspond to gaps in

201 settlement distribution, but not all gaps in settlement distributionsuch as rock outcrops correspond to gaps in baobab distribution. Large baobabs and small baobabs show attraction over all distances analyzed (Figure 7, p. 222). However, this attraction is significant only at distances less than c.300 m, and only for two of the three rectangular sub-regions. Observed significance is marginal in all cases. Baobab-settlement spatiality. Baobabs and settlements are attracted at nearly all spatial scales for all pairs of data sets examined (Figure 8, p. 224). However, observed attraction is significant only at distances less than c.500 m. Strongly significant attraction is observed in the distribution of all baobabs in relation to all settlements. Decreasingly significant attraction is evident in the distribution of: a) large baobabs and old settlements; b) small baobabs and young settlements; and c) small baobabs and old settlements. Finally, the locations of large baobabs and young settlements displayed essentially no significant attraction; in the Rectangle 3 subregion, attraction between these data sets shows minimal significance. The statistical significance of baobab-settlement attraction is, for most pairs of data sets, considerably stronger than that observed for attraction between large and small baobabs (Figs. 7, 8). Finally, spatial relationships between baobabs and settlements vary between rectangular sub-regions (Figure 8, p. 224). In some portions of the study area, particularly Rectangle 2, baobabs and settlements display repulsion over intermediate distances. For large baobabs and young settlements, this repulsion is marginally significant at a distance of about 2 km. Discussion Baobab habitat. Baobabs occur in a wide range of habitats in the research area, but abundance and reproductive success are not uniform across this range (Table 1, p. 226). Baobabs are most abundant in occupied and abandoned settlements and fields. The tree is also

202 relatively abundant on cliffs and steep slopes (Figure 6, p. 220), even though previous authors have made only passing mention that it occurs in such habitat (Mullin 1992: 66; Wickens 1982: 188). Of course, a conclusion based primarily on observation of large individuals would be that baobabs are rare on cliffs and steep slopes: large individuals are significantly underabundant in such habitats (Table 1, p. 226). Cliffs and steep slopes are geomorphically unstable, and baobab mortality due to substrate collapse and rock fall appears to be high and positively correlated to tree size (cf. Larson et al. 2000). Small trees can be difficult to see in many situations, but time spent searching increases the rate at which they may be found (Figure 4, p. 218). Indeed, small baobabs are frequently encountered along cliffs and steep slopes, and cliff-side gallery forest hosts nearly as many small baobabs as occupied settlements and fields (Table 1, p. 226). Baobab abundance in settlement and field sites and the overabundance of small baobabs along cliffs and steep slopes indicates: a) the importance of primates, including humans, as seed dispersers; b) the trees adaptation to dry soil conditions; and c) the possible intolerance of small baobabs to fire. Non-human primates disperse baobab seeds (Wickens 1982), and baboons and chimpanzees may account for the trees abundance on rock outcrops. In the fruiting season (November-January), baboons are frequently observed eating fallen baobab fruit, and frequently travel along and rest on cliffs (Duvall, unpublished data). Chimpanzees and their nests are most frequently encountered near large, fruiting baobabs and along cliffs from November-January (e.g. Moore 1985), and baobab seeds are amongst the most common food remains in fecal samples from this period (see Chapter 6). While other animals certainly disperse baobab seeds, no other large animals appear to share spatial ecology as closely with the baobab. The distribution of human settlement also corresponds strongly to baobab distribution, as discussed

203 below. Other human activities, such as cultivation in fields distant from settlements and fruit consumption while traveling along paths, certainly account for the presence of some baobabs away from settlement sites, but nearly all areas along cliffs and steep slopes are considered uncultivable (see Chapter 3) and are nearly inaccessible to humans. Human activities cannot reasonably account for the abundance of baobabs along cliffs and steep slopes. Cliffs and steep slopes provide a wide range of habitats in terms of soil moisture (Larson et al. 2000). Ledges and soil pockets amongst bouldersthe microhabitats most frequently occupied by baobabs (Figure 6, p. 220)are relatively xeric. While most baobabs along cliffs and steep slopes occur in cliff-side gallery forest vegetation, which has a high abundance of mesophytes (Duvall 2001), baobabs do not occur in moist microhabitats. Additionally, only 2.2% of observed individuals occur along drainage channels, though only in drier portions of channels without riparian gallery forest vegetation (Table 1, p. 226). Settlement sites, where most small baobabs occur, also seem to represent relatively xeric habitat; dryness is a primary criterion in Maninka site selection. Vegetative cover is relatively low in settlements, meaning that insolation and run-off are relatively high (Belsky et al. 1993). As an indication of the dryness of settlements as tree habitat, several woody xerophytesparticularly Bauhinia rufescens Lam., Calotropis procera (Aiton) W.T. Aiton, and the domesticated date palm (Phoenix dactyllifera L.)occur only in settlements in the study area. Fenner (1980) shows that baobab is a drought-adapted species, but none have considered how its physiological ecology may relate to its association with settlements. Although better knowledge of the characteristics of settlements as tree habitat is necessary, these results suggests that baobab-habitat association occurs not simply because humans disperse seeds to settlements, but also because settlement sites are suitably dry for baobabs. Other wild fruit trees with dry

204 habitat preferencessuch as Tamarindus (Arbonnier 2000) and Borassus (which requires a high water table: Sambou et al. 1992)also increase in abundance at settlement sites during site occupation, according to ethnographic data and vegetation analysis (Duvall in review a). In contrast, wild fruit trees that are associated with mesic habitatssuch as Spondias mombin and Cordia myxa L. (Arbonnier 2000; Geerling 1982)generally increase in abundance only after site abandonment, once trees surviving human occupation have grown large enough to create cooler and moister soil conditions (Belsky et al. 1993) and attract non-human frugivores. While humans may contribute to the dryness of settlement sites through vegetation clearing and earthworks, non-anthropogenic physical processes are probably the primary cause of the dry soil conditions that make settlement sites suitable for baobabs. Additionally, cliffs and settlements are both well protected from fire. Wildfires, especially intense ones, are unable to reach most portions of cliffs and steep slopes (Larson et al. 2000). Fires lit in and around settlements and in actively cultivated fields are closely controlled; there is generally insufficient fuel on the ground in settlements or fields to sustain uncontrolled fires. There has been no study of baobabs fire tolerance, although based on general firevegetation relationships several authors have concluded that baobabs, especially small ones, are poorly adapted to frequent or intense fires (Esenowo 1991; Napier-Bax & Sheldrick 1963; Palmer & Pitman 1972; Simpson 1995; Wilson 1988). While baobab has relatively thin bark, suggesting vulnerability to fire, it can sprout from its roots and regenerate cambium (Wickens 1982), indicating a capacity to recover from injury. Indeed, Baumer (1983: 67) states that [t]he tree is not much affected by bush fires. Small baobabs are overabundant in occupied settlements, occupied fields, and along cliffs, and underabundant in old fields and woodland vegetation. In contrast, large baobabs are overabundant in old fields and woodland, habitats that

205 are generally not protected from fire (Laris 2002; Mbow et al. 2000). These results suggest that ecological conditions in fire-controlled habitats are optimal for young baobabs, while conditions in fire-prone habitats are tolerable to large, but less so to small, individuals. Fire protection may be a significant aspect of settlements as tree habitat that contributes to baobab-settlement attraction. Human-baobab biogeography. Settlements and baobabs are attracted at most spatial scales (Figure 7, p. 222), but there is no single reason for the observed attraction. A portion of the attractionespecially for larger spatial scalesis due to the fact that the preferred habitat for human settlement is the same as that in which baobabs are most abundant: areas with deep, arable soil, dominated by woodland vegetation. More specifically, Maninka knowledge of baobab-human interaction indicates that human settlement leads to the development of baobab groves at settlement sites, and that settlements are rarely established under preexisting baobabs. Point pattern and chi-square analyses support Maninka knowledge. In particular, large baobabs and young settlements are the only point sets that do not exhibit significant attraction: young settlements are rarely established under preexisting, large baobabs. All other point sets, including small baobabs and young settlements, display significant attraction at short distances: baobab recruitment occurs significantly more than expected at settlement sites. Additionally, point pattern analysis of the distributions of large and small baobabs (Figure 7, p. 222) shows that small baobabs are more significantly associated with settlements than with large baobabs. While non-anthropogenic baobab recruitment certainly contributes to the significant attraction between small baobabs and old settlements, settlement-baobab attraction is not simply the result of natural recruitment associated with large individuals.

206 Repulsion observed between baobabs and settlements at some scales also suggests dependence, although baobab-settlement repulsion is but marginally significant, only at distances of c.2 km, and only in the Rectangle 2 sub-region (Figure 7, p. 222). New settlements are most frequently established by young men seeking access to more arable land than available to them in their home village. Men establishing settlements often seek sites that are several kilometers from preexisting, occupied or abandoned settlements in order to avoid conflict over historically established land or tree usufruct. This aspect of settlement practice creates observable repulsion between young settlements and baobabs that parallels repulsion between young and old settlements. African farmers have profoundly altered plant distributions through migration, settlement, and cultivation (Maranz & Wiesman 2003; O'Brien & Peters 1998). Baobab groves created at settlement sites in semi-arid Mali are analogous to forest islands farmers create at settlement sites in sub-humid Guinea (Fairhead & Leach 1996), even though the two vegetation features are structurally and compositionally different. Although Fairhead and Leach (1996) describe a teleological process of vegetation enrichment, the creation of baobab groves at settlement sites in southwestern Mali is an unintentional outcome of settlementalthough farmers clearly recognize, understand, and anticipate this outcome when establishing new settlements. While baobab fruit collection contributes most visibly to baobab-settlement attraction, other aspects of settlement practiceparticularly site selection and fire protection are also important. Past assessments of human-baobab interactions have relied largely on qualitative analyses of incomplete censuses of baobabs and settlements in focal landscapes. In many cases, impressions of landscape characteristics have been the primary data used to support the

207 conclusion that baobab and settlement distribution are spatially dependant (Aubrville 1950; Chevalier 1906; Hobley 1922; Sikes 1972; Taylor 1960). Humans have modified woodland composition throughout Africa (e.g. Ballouche & Neumann 1995; Boffa 1999; Maranz & Wiesman 2003; O'Brien & Peters 1998; Pullan 1974), but it is vital to identify clearly how humans alter vegetation characteristics to avoid inaccurate or oversimplified representations of human-environment interactions that support socially unjust and ecologically short-sighted policies (Bassett & Koli Bi 2000; Leach & Mearns 1996; Turner 2000). Human settlement increases baobab recruitment in parts of the landscape where small baobabs are otherwise uncommon (i.e. flat areas dominated by woodland vegetation), but this influence is significant only over distances <c.500 m. Ecological factors other than settlementsuch as seed dispersal by non-human primates, and other human activitiesmust account for the occurrence of baobabs in many parts of the landscape, especially along cliffs and steep slopes. At a scale of tens of kilometers, baobab distribution cannot be attributed entirely to humans, although the trees dispersion and abundance would be lower in the absence of human settlement. Based on data collected across West Africa, Maranz and Wiesman (2003) propose that economically important, semi-domesticated trees in African agroecosystems reproduce independently of human activities, but humans alter gene frequencies by culling self-sown trees with unfavorable phenotypes. The observed, landscape-scale distribution of baobab is consistent with Maranz and Wiesmans model, although their model may underestimate the dependence of tree reproduction on human activities. Specifically, the overabundance of small baobabs in cliff habitats indicates that the tree does reproduce independently of humans, but the underabundance of large (i.e. reproductive) trees along cliffs, along with the overabundance of large trees in abandoned settlements and fields, shows that the baobab population depends on human activities

208 to maintain a recruitment rate sufficient for population replacement. If baobab reproduction was predominantly independent of human activities, the tree would be much rarer because few trees would grow large enough to fruit. However, multi-scale analyses are necessary to understand more completely how humans affect the reproduction and distribution of economically important, African trees. For instance, landscape-scale study of the distribution of Vitellaria paradoxa is necessary to determine how this tree reproduces in landscapes where it has been introduced in the past 200 years (cf. Kelly et al. 2004; Maranz & Wiesman 2003), while regionto continent-scale genetic analysis is needed to determine whether humans introduced baobab in the present study area. The present results suggest concretely how land-use changea factor implicated in poor recruitment for many tree populations in semi-arid Africa (Assogbadjo et al. 2005; Lykke 1998; Sambou et al. 1992; Schreckenberg 1999; Wilson 1988)may negatively affect baobab reproduction. Shifting cultivation and settlement, including the establishment and abandonment of fields and settlements, has been practiced in semi-arid Africa for centuries (Raynaut 1997; Richards 1978; Stone 1996). Parkland vegetation, dominated by valued wild fruit trees, is clearly a long-term result of agroforestry practices that increase the relative abundance of highly valued trees in farmed and settled parts of landscapes (Boffa 1999; Maranz & Wiesman 2003; Pullan 1974; Schreckenberg 1999). Parkland vegetation does not occur throughout semi-arid Africa, but the distribution and abundance of highly valued trees in areas without parkland vegetation also results from long-term management practices. Most reported cases of poor recruitment for highly valued wild trees are from areas where land-use intensification is proceedingincluding the expansion of cultivated area paired with the reduction in fallow land, and increased commodification of non-timber forest productsoften at the expense of land-

209 extensive practices (Assogbadjo et al. 2005; Johansson 1999; Lykke 1998; Sambou et al. 1992; Schreckenberg 1999; Wilson 1988). In the research area, baobab population replacement rate is linked to the rate of settlement establishment and abandonment: small baobabs are most abundant in occupied settlements and fields, and large (i.e. reproductive) baobabs are most abundant in abandoned settlements and fields (Table 1, p. 226). Processes that reduce the rate of settlement establishment or abandonment will negatively affect recruitment rates of tree species associated with settlements. Conservation policies that prohibit shifting cultivation or settlementsuch as those recently enacted in the present research area (see Chapter 2)represent non-economic processes that reduce rates of settlement establishment and abandonment. Thus, such policies may unintentionally reduce recruitment or dispersion of wild fruit trees associated with settlements. While such policies are unlikely to lead to extinction or local extirpations of effected trees and may be necessary to address short-term threats to biodiversity resources, declines in wild fruit tree populations caused by land-use change may have unforeseen consequences for ecosystems over longer terms if effected tree species are important resources for wildlife (cf. Fimbel 1994a; Fimbel 1994b; Wilkie & Finn 1990). Conclusion The results show that baobab population regeneration and human settlement establishment are spatially dependent. Human settlements are habitat patches that are suitable for baobabs, and baobabs adaptation to human fruit preferences means that the tree has gained a reliable dispersal agent that allows it to colonize these habitat patches. The trees success in these habitat patches, long recognized through observation of baobab groves at settlement sites, is not simply a function of seed accumulation at settlement sites. Other aspects of settlement

210 practiceespecially the selection of dry, well drained sites with deep soil, and the prevention of fires in settlementsmake settlement sites suitable habitat for baobabs. Baobabs are also relatively abundant in similarly dry and fire-protected habitats on cliffs and steep slopes, although these sites are quite unlike settlements in other ways. The trees abundance on cliffs also indicates its adaptation to baboon and chimpanzee feeding behavior. Indeed, these animals may, over the long term, benefit from increased baobab dispersion and abundance resulting from human settlement. Other highly valued, wild fruit trees likely have similar relationships with human settlement and non-human frugivores, and the persistence of these relationships is linked to long-term patterns of land use.

211 Figures and tables for Chapter Four

212 Figure 1. Baobab range and research area. Map 1: Africa, showing location of Map 2 and baobab distribution, which is redrawn and simplified from Wickens (1982). Distribution on islands (except Madagascar) not shown. Map 2: Western Mali, showing location of research area. Map 3: Research area, showing location of Solo, all occupied settlements, and cliffs in the research area.


214 Figure 2. Baobab and settlement data maps. Abbreviations: B=baobab/s, Q=quintile/s, S=settlement/s. Dashed rectangles indicate the rectangular sub-areas used to test significance in point pattern analysis. These sub-areas are referred to by number, shown in a box near the lower, left-hand corner of each sub-area. Panel 1: Bullet points represent all baobab individuals; grey circles represent circular areas (500 m diameter) centered on all settlement sites. This diameter is chosen to provide cartographic representation of the distance at which baobab-settlement attraction is significant in most cases (see text and Figure 8, p. 224). Panel 2: Triangles represent all settlements, with shading indicating quintile membership of each settlement. Panels 3-6: Bullet points and gray circles as described for Panel 1. Smallest baobabs and Youngest settlements depict only quintiles 1 and 2 of the baobab and settlement data sets. Largest baobabs and Oldest settlements depict quintiles 4 and 5 of these data sets. These panels represent to the paired point sets described in Table 2 (p. 227), whose observed L-function values are shown in Figure 8 (p. 224).


1 3 All baobabs 2
Latitudinal distance (meters)

1 3 2

SQ 1&2 SQ 3 SQ 4&5

1 3 Smallest B, Youngest S

1 3 Largest B, Youngest S

1 3 Smallest B, Oldest S

1 3 Largest B, Oldest S

Longitudinal distance (meters)

216 Figure 3. Photos of baobabs in settlements. Photo (a): What appear to be pebbles embedded in the ground surface are often baobab seeds. Arrows indicate three seeds in this photo. Photo (b): During the rainy season, hundreds of baobab seeds germinate in rubbish mounds, but most die during the dry season. Photo (c): The few seedlings that survive happen to grow in wellprotected situations, such as under granaries. Arrows indicate three saplings emerging from under a granary that had been built about seven years earlier. These saplings are stunted as a result of frequent leaf collection and browsing livestock. Photo (d): Baobabs that outlast human occupation of a settlement site often thrive to dominate vegetation in abandoned settlements. Arrows indicate three medium-sized baobabs that are nurse trees for various other wild fruit trees.



Figure 4. DBH measurement in relation to research effort. Trend line is a linear regression of DBH measurements for observed baobabs, arranged in order of date encountered.

219 Figure 5. Baobab size-class distribution. Size classes are 10-cm DBH intervals.

220 Figure 6. Photos of baobabs in cliff habitats. Photo 1: Most baobabs on cliffs grow on narrow ledges, which baboons and chimpanzees use in traveling along rock outcrops. This photo, taken from the top of a cliff looking down at a ledge about 25 cm wide and 2 m below the cliff top, shows a baobab sapling (its base is to the right of the arrow) that has partially fallen, turning over the soil mat in which it is rooted. (The branches to the right are the crowns of trees rooted at the base of the c.20-m cliff.) Baobab mortality on cliffs appears to be very high due to the geomorphic instability of this habitat. Photo 2: Baobabs that attain large size on cliffs are those that happen to grow in relatively flat, rocky patches of soil trapped between boulders. Both of these cliff microhabitats experience xeric soil moisture conditions although they may be in or adjacent to mesophytic cliff-side gallery forest. These two medium-sized trees have annular scars near their bases that indicate humans harvested their bark for rope when the trees were smaller.


222 Figure 7. Large baobab-small baobab bivariate point-pattern analyses. Observed L-function values depicted with solid lines; dashed lines depict simulation envelopes derived from 99 random toroidal shifts of data sets. Values >0 indicate spatial attraction between large and small baobabs; values <0 indicate repulsion. Where observed values lie outside simulation envelopes, observed attraction/repulsion is significant (p<0.01).



Figure 8. Baobab-settlement bivariate point-pattern analyses. Observed L-function values depicted with solid lines; dashed lines depict simulation envelopes derived from 99 random toroidal shifts of data sets. Values >0 indicate spatial attraction between baobabs and settlements; values <0 indicate repulsion. Where observed values lie outside simulation envelopes, observed attraction/repulsion is significant (p<0.01).


226 Table 1. Baobab-habitat associations. Symbols used to indicate statistical significance:


=overabundant, p<0.001; ++=overabundant, p<0.01; +=overabundant, p<0.1;

=underabundant, p<0.001; =underabundant, p<0.01; and =underabundant, p<0.1. Abbreviations: aban.=abandoned; indivs.=individuals; occ.=occupied; Q=quintile. Habitat for each baobab was characterized by selecting one category (e.g. pathside, slope >36, woodland) per environmental factor (i.e. land use, topography, vegetation). The categories Abandoned settlement and Occupied settlement include only trees found 100 m from the center of a settlement site or 20 m from huts found 100 m from a settlement center. Other land-use categories based on observation of current use, or oral historical evidence for past use. For quintile DBH ranges, see Table 2 (p. 227). All indivs. 1240 443 (35.7%) 336 (27.1%) 331 (26.7%) 76 (6.1%) 37 (3.0%) 17 (1.4%) 881 (71.0%) 227 (18.3%) 105 (8.5%) 27 (2.2%) 918 (74.0%) 122 (9.8%) 116 (9.4%) 61 (4.9%) 23 (1.9%) Q1 248 80 53 59 35+++ 20+++ 1 181 23 40+++ 4 142 57+++ 45+++ 3 1 Q2 248 89 72 62 18 3++ 4 178 34 27 9 192 22 25 3 6 Q3 248 107++ 46 80+ 9 4 2 165 53 22 8 179 15 23 21+ 10+ Q4 248 81 74 74 9 6 4 174 61++ 10 3 197+ 18 10 20+ 3 Q5 248 86 91+++ 56 5 4 6 183 56+ 6 3 208+++ 10 13 14 3

Number of trees Land use Aban. settlement Aban. field No history of past use Occ. settlement Occ. field Pathside Topography Slope <5 Slope 5-36 Slope >36 Drainage channel Vegetation Woodland Occ. settlement or field Cliff-side gallery forest Bamboo thicket Rupicolous scrub

227 Table 2. Baobab and settlement quintile characteristics, and paired data sets used in pointpattern analyses. Abbreviations: B=baobab/s; DBH=diameter at breast height; Q=quintile; S=settlement/s. Section 1: Second and third columns give ranges per quintile of the quantitative variable determining quintile membership (DBH for baobabs, years since establishment for settlements). Fourth column gives number of settlements per quintile; membership per quintile unequal because several settlements have same estimated ages. All baobab quintiles have 248 members. Section 2: Bivariate point-pattern analyses were conducted as described in the text for each point pair indicated, in the study area as a whole and in three rectangular sub-regions (Figure 3, p. 216). Section 1: Quintile characteristics Q B DBH S Age S per (cm) (yrs) Q 1 1-19.8 4-24 19 2 3 4 5 20.060.0 60.5124.3 124.5193.3 195.0455.0 29-45 47-66 69-110 120250 17 19 16 18 Section 2: Paired point sets used in bivariate analyses. Point Ecological meaning sets All B, Are baobabs associated with All S settlements? BQ 1&2, Are small baobabs associated with SQ 1&2 young settlements? BQ 1&2, Are small baobabs associated with old SQ 4&5 settlements? BQ 4&5, Are large baobabs associated with old SQ 4&5 settlements? BQ 4&5, Are large baobabs associated with SQ 1&2 young settlements? BQ 1&2, Are small baobabs associated with large BQ 4&5 baobabs?

228 Chapter Six: Chimpanzee diet, habitat use, and human settlement in Mali

Abstract Humans have profoundly altered vegetation composition in Africa through millennia of settlement and cultivation. In many areas, a long-term effect of settlement has been the development of vegetation patches at settlement sites, where structure and composition differ from that in surrounding vegetation. These patches represent potentially valuable habitat for chimpanzees and other frugivores, because settlement increases the relative abundance of wild fruit trees at settlement sites. In southwestern Mali, settlement creates baobab groves at settlement sites, and past observations of chimpanzee behavior suggest that chimpanzees frequently forage in baobab groves, at least seasonally. The purpose of the present paper is to determine spatio-temporal characteristics of chimpanzee habitat use in Mali, and particularly to determine if the animal significantly uses abandoned settlements as habitat. The results show that chimpanzees use abandoned settlements especially during the time of year when baobab fruit is an important dietary component. During other times of the year, forest habitats along sandstone outcrops are more heavily used, because permanent water sources and food patches are more abundant along outcrops. Chimpanzees are ecologically linked to groundwater stored in sandstone outcrops, like other rare and biogeographically noteworthy species in southwestern Mali. Anthropogenic baobab groves increase the abundance and distribution of chimpanzee food patches, and appear to be especially important during times of the year when fruits produced by plants characteristic of cliff habitats are not abundant. Recognizing indirect human-chimpanzee interactions is necessary for reducing unforeseen, long-term consequences of conservation policies.

229 Key words chimpanzees; agriculture; conservation; Mali; ecological history

Introduction In West Africas semi-arid, Sudanian woodland zone, human settlement has been characterized historically as destructive of natural resources, especially woody vegetation (Aubrville 1949a; Chevalier 1938; Keay 1959). The discourse of deforestation and desertification has arisen, in part, from the view that rural settlements cause declines in vegetation density, productivity, and diversity (Bassett & Crummey 2003; Fairhead & Leach 1996; Fairhead & Leach 1998). While urbanization has led to loss of woody cover around cities (Becker 2001), research on vegetation ecology in rural areas has shown that although the effects of settlement are variable, settlements are generally not deforested patches. In some areas, where land-use intensification is proceeding, the density or diversity of woody plants has decreased, or has the potential to decrease (Lykke 1998; Nyerges 1989; Schreckenberg 1999). However, there are also examples of afforestation associated with rural settlement: in Guinea and Ghana, settlements become forest islands in a savanna matrix due to purposeful management of settlement sites (Amanor 1994; Fairhead & Leach 1996); in Cte dIvoire and Cameroon the density and abundance of woody species is increasing in woodlands around settlements as an unintentional outcome of increased livestock ownership (Bassett & Boutrais 2000; Bassett & Koli Bi 2000); and in Mali, human use of wild fruits and management of settlement sites directly and indirectly create patches of wild fruit trees in woodlands (see Chapters 5 and 6). Such longterm, often subtle changes in vegetation structure and composition are indicative of the

230 profoundly anthropogenic character of many types of African vegetation (Lovett & Wasser 1993; Lovett & Haq 2000; Maranz & Wiesman 2003; O'Brien & Peters 1998). What effects do gradual, long-term, anthropogenic changes in vegetation characteristics have on wildlife? The response of wild animals to disturbance depends on the type and degree of change, as well as a species behavioral ecology and body size (Johns & Skorupa 1987; Plumptre 2001; Plumptre & Reynolds 1994). Drastic, rapid vegetation changes, such as nonselective logging in rainforest areas, decimate most wildlife populations associated with the original habitat. Indeed, the drastic land-cover changes that have occurred globally in the past century have made habitat loss the main threat to plant and animal populations worldwide (Hilton-Taylor 2000; Walter & Gillett 1998). This global, relatively recent situation does not describe all cases of vegetation change caused by human disturbance, especially those that have occurred slowly, over hundreds of years. Over the course of centuries to millennia, human management of occupied and abandoned settlement, field, and garden sites has altered vegetation composition throughout the tropics (Bale 1994; Croll & Parkin 1992; Denevan & Padoch 1988; Fairhead & Leach 1996; Janzen 1998). One aspect of this change has been the development of vegetation patches that are attractive to wildlife for various reasons, but primarily as food patches (Medellin & Equihua 1998; Naughton-Treves 2002; Thiollay 1995; Thomas 1991). These food patches may significantly enhance habitat quality for some wildlife species in human-occupied landscapes, and reflect long-term co-adaptation between humans and wildlife (Bahuchet & Garine 1990; Fimbel 1994a; Gadgil et al. 1993; Greenberg 1992). Their significance may be muted, however, by hunting: farmers often depend on abandoned settlement, field, and garden sites to produce

231 meat (Bahuchet & Garine 1990; Denevan et al. 1984; Fairhead & Leach 1996; Naughton-Treves 2002; Nietschmann 1973). Wildlife response to long-term, anthropogenic vegetation change has not been studied in the semi-arid tropics, where settlement and agricultural practice, and the composition and ecology of plant and animal communities, differs strongly from rainforests. Additionally, studies of wildlife use of abandoned settlement, field, and garden sites have often focused on animals valued for meat (although see: Fimbel 1994a; Fimbel 1994b), and frequently do not contextualize wildlife activities in humanized sites by providing comparable data for animal behavior across entire landscapes, including non-humanized portions. The present paper looks specifically at how chimpanzee (Pan troglodytes verus) abundance varies across a Malian landscape in which human settlement over the last c.400 years has significantly changed the distribution and composition of certain plant communities. In African rainforests, chimpanzee populations have almost uniformly declined due to short-term vegetation change, especially clearing for logging or agriculture (Chapman & Lambert 2000; Kormos et al. 2003; Plumptre 2001; Plumptre & Reynolds 1994). However, in rainforest settings where agricultural disturbance results in the development of persistent patches of woody plants with edible fruitsand where hunters do not purposefully target chimpanzees chimpanzee abundance has not declined, and the animals use these disturbed patches more frequently than expected (Fimbel 1994a; Fimbel 1994b; Wilkie & Finn 1990). Circumstantial evidence suggests that chimpanzees in Mali also rely on vegetation patches at abandoned settlement sites. Specifically, Moore (1985: 60), after surveying chimpanzee distribution in southwestern Mali, found that many of the chimp nests we observed occurred near fruiting baobabs. Baobabs (Adansonia digitata L.) dominate vegetation at

232 abandoned settlement sites in many parts of Africa (Wickens 1982), and in southwestern Mali baobab groves develop at settlement sites directly and indirectly as a result of human activities (see Chapter 5). However, other findings seem to conflict with this circumstantial evidence. In particular, humans settle and farm relatively flat areas with deep soil in southwestern Mali (see Chapters 2 and 4), while chimpanzee abundance is strongly associated with cliffs and steep slopes (Granier & Martinez 2004; Moore 1985; Pruetz et al. 2002). The purpose of the present paper is to determine spatio-temporal characteristics of chimpanzee habitat use in Mali, and particularly to determine if the animal significantly uses abandoned settlements as habitat. This paper analyzes a range of data, from: 1) systematic surveys of chimpanzee abundance, 2) studies of chimpanzee nesting and feeding ecology, 3) maps of the distribution of surface water and chimpanzee food-plant patches, and 4) ethnographic interviews on local settlement history and chimpanzee behavior. These disparate but complementary information sources support the view that human settlement creates valuable habitat patches for chimpanzees in semi-arid West Africa. Human-chimpanzee interactions in semi-arid West Africa The northernmost limit of chimpanzee distribution occurs near 13 N in Senegal and Mali (Butynski 2003). Chimpanzees have been known scientifically in this area for nearly 30 years, and various researchers have contributed to a relatively small, largely exploratory literature on Malis chimpanzees (Duvall 2000; Gagneux et al. 1999; Granier & Martinez 2004; Moore 1985; Pavy 1993; Sayer 1977). Chimpanzee ecology in Mali is probably quite similar to that in southeastern Senegal, where research on the apes behavioral ecology has been more abundant and substantial (Baldwin et al. 1982; Baldwin et al. 1981; Bermejo et al. 1989; Marchant & McGrew 2005; McBeath & McGrew 1982; McGrew et al. 1981; McGrew et al. 1988; Pruetz et

233 al. 2002). Many researchers have found that chimpanzee abundance is strongly associated with topography; nests (which chimpanzees build nightly for sleeping) are from eight (Granier & Martinez 2004) to 800 (Pruetz et al. 2002) times more abundant in forests along cliffs and steep slopes than in relatively flat, woodland areas. However, around Mt. Assirik, Senegal, observers have recorded more nests in relatively flat, woodland areas rather than in forests associated with abrupt topography (Baldwin et al. 1982; McGrew et al. 1981). Interactions between chimpanzees and humans in semi-arid Africa have been poorly studied, although three interactions that represent threats to chimpanzees are certain in Senegal and Mali. First, humans rarely hunt chimpanzees (Carter et al. 2003; Duvall et al. 2003; Granier & Martinez 2004), because most people consider chimpanzee flesh inedible as food. Limited hunting exists because some people consume chimpanzee meat for medicinal use (Duvall & Smith 2005), and there is a small but persistent pet trade sustained by hunting (Carter et al. 2003; Duvall et al. 2003; Granier & Martinez 2004). Due to the animals relatively low population size in these countries (up to c.1500 individuals in Mali: Duvall et al. 2003) and low reproductive rate, even a very low rate of hunting could pose a serious threat to the population (Moore 1985). Although all sources indicate a low absolute level of hunting, this threat remains a crucial issue for chimpanzee conservation, and better knowledge of chimpanzee population size and the rate of human hunting of chimpanzees is needed to manage this threat successfully. Second, humans and chimpanzees compete for wild plant resources, especially fruits. In Senegal, commercial harvesting of wild fruits for sale in cities has reduced local abundance of chimpanzee food during certain seasons, leading chimpanzees to consume lower-quality foods (Carter et al. 2003; Pruetz 2002). Finally, road building, agricultural expansion, and industrial mining all cause habitat loss at an unknown rate (Carter et al. 2003; Duvall et al. 2003). In particular,

234 conservationists are concerned that indigenous settlement practices in Malis Bafing Biosphere Reserveespecially the establishment and abandonment of dispersed farming hamlets represent the expansion of settlement into previously unoccupied areas of wildlife habitat (see Chapter 2). While there is no frontier-style settlement expansion occurring in the Bafing area (Chapter 2), the effects of shifting settlement on chimpanzee habitat and behavior are unknown. Research area Research occurred in an area of 183 km2 around the village of Solo (1258' N, 1026' W) in Malis Bafing Biosphere Reserve (Figure 1, p. 258). In this area, topographic complexity associated with bedrock outcrops helps create a wide range microenvironments (Jaeger 1959; Lawesson 1995). In the northern portion of the research area, sandstone plateaus dominate the landscape, rising 200-300 m above surrounding lowlands (IGM 2001). The southern portion is relatively flat, with few bedrock outcrops, all of which are dolomite, not sandstone. Erosion of the sandstone plateaus has formed narrow ravines, rocky slopes, and plains with relatively infertile sandy and silty soils. Dolomite outcrops erode more uniformly to form rounded inselbergs surrounded by silty soil. Ferricrete hardpans and bare bedrock surfaces, with very dry soil conditions, are common throughout the area (Dames & Moore 1992; Jaeger & Jarovoy 1952; Michel 1973). Groundwater seeps to the surface permanently or seasonally in some locations where sedimentary layers in the sandstone have been exposed (DCTD 1990). If seepage occurs in topographically sheltered locations, very humid microclimates exist (Duvall 2001). Elsewhere, permanently moist habitats are uncommon, and are primarily deep depressions in seasonal streambeds. Soils in most locations are driest from April to June, when air temperature and potential evapotranspiration peak (FAO 1984).

235 Seasonal changes in rainfall and potential evapotranspiration control plant phenology and the availability of surface water. Precipitation is highly seasonal and averages about 1100 mm per year, with high interannual variation (FAO 1984; Leroux 2001). Most rain falls during JuneSeptember, but brief, intense rainstorms also occur during April-June; rainfall is minimal during November-March (Dames & Moore 1992; FAO 1984; Leroux 2001). Woody plant phenology varies between habitats based on the dominant reproductive strategy in each habitat, largely a function of annual soil moisture variation (Breman & Kessler 1995; de Bie et al. 1998; Rathcke & Lacey 1985). Many deciduous species (dominant in woodland areas but also present in forests) rely on stored energy to produce fruit during April-June or during one other period, while many evergreen species (dominant in forests) flower or fruit throughout the year (Arbonnier 2000; Breman & Kessler 1995; de Bie et al. 1998). Woodland vegetation dominates most of this landscape, especially in areas with relatively deep, fertile soil. Forest patches occur in topographically protected microhabitats with moist soil conditions along bedrock outcrops. These patches are important chimpanzee habitat (Duvall 2000; Granier & Martinez 2004; Moore 1985; Pavy 1993). Locations with shallow or infertile soil host patches of edaphic bushland or grassland. Based on woody species composition, fifteen vegetation types have been described for the area (Table 1, p. 264). Maninka farmers, who have occupied this landscape for at least 400 years (Samak et al. 1986), have altered vegetation characteristics. Most sites with arable soil have been farmed in the past, and all sites have been subject to at least low intensity or frequency disturbance. Designating different types of vegetation primary or secondary based on past human disturbance is not helpful in understanding vegetation ecology, because the effects of human activities on vegetation are variable, and many types of undisturbed vegetation are

236 compositionally and structurally similar to disturbed vegetation (see Chapter 4). The Maninka settle and cultivate sites with deep, arable soil and good drainage. Large areas of the landscape have never been cleared for agriculture or settlement: rocky areas, plateau tops, and sites with poor soil or drainage are used only for seasonal livestock grazing, wild plant and honey collection, and hunting (Duvall 2001; Samak et al. 1987). Many abandoned human settlement sites host distinctive vegetation dominated by disturbance-adapted shrubs and by trees with edible fruits (see Chapters 5 and 6). In particular, past settlement has led directly and indirectly to the development of many baobab groves across the landscape (see Chapter 5). Data collection and analysis Five types of data were used: 1) observations of chimpanzees and night nests; 2) contents of chimpanzee fecal samples; 3) samples of vegetation composition; 4) observations of plant phenology; 5) mapping of permanent surface water sources and chimpanzee food-plant patches; and 6) ethnographic interviews of indigenous farmers and hunters. Chimpanzee and nest observations. During May-July 2003, reconnaissance surveys and ethnographic interviews of indigenous hunters revealed approximately 20 sitessuch as forest patches and baobab grovesthat chimpanzees appeared to use frequently for nesting or foraging. During January-December 2004, the researcher and six research assistants visited each of these sites twice weekly for 38 weeks (observations were made on 241 total days) to record observations of chimpanzees and their night nests. Specifically, each research assistant walked one of six survey loops (Figure 2, p. 259) that passed through several of the sites that chimpanzees appeared to use frequently. However, most of the length of each survey loop was in habitatssuch as wooded grasslandwhere evidence for chimpanzee abundance is low. Repeated visits to the sites identified in 2003 revealed that chimpanzees did not frequently use

237 many of them. Nonetheless, the layout of survey loops was not modified during 2004, in order to collect systematic data on variation in chimpanzee abundance, and because increased familiarity with the research area during 2004 did not reveal any additional sites that chimpanzee appeared to use frequently. Two types of observations were recorded when walking survey loops: 1) visual observations of chimpanzees, and 2) nests constructed since the previous survey. Locations and times of all observations were determined using Garmin 12-XL GPS units. For each chimpanzee sighting, the initial activity of observed chimpanzees was recorded, if possible, and observation sites were searched for feces and evidence of feeding. Observers did not attempt to follow observed chimpanzees. Multiple observations of a single group may have been recorded as single observations of multiple groups, but this possibility is unlikely. Only 3 of 48 observations followed an earlier observation on a single survey loop on one day, and several kilometers separated these same-day observations. Similarly, different survey loops that were walked on the same day were walked at approximately the same time (approximately 8:00 a.m.-2:00 p.m.) and were separated by 5-20 km; examination of observation times shows that chimpanzee groups seen on different survey loops on one day could not have been a single group. To locate nests, observers walked at c.4-6 km/h, searching from side to side for nests visible from the survey loop. Upon observing a nest, the observer left the survey loop to examine the nest and search for other contemporaneous nests belonging to the same nest group. The number of nests per nest group was counted. Nest groups were defined as all contemporaneous nests within 20 m of one another (cf. Blom et al. 2001). In practice, nearly all nest groups were clearly discrete, isolated by several hundred meters from contemporaneous

238 nests. Contemporaneous nests were defined as those that had been constructed since the previous survey; the period between surveys per loop was 4-7 days. These nests were considered fresh because their leaves remained green, flexible, and not dry, and their construction could be attributed to the preceding 4-7 days. Data collection began on 31 January 2004, ceased from 16 July to 30 August 2004, and continued until 15 December 2004; for the weeks following 31 January and 30 August, the initial survey of each loop served to identify all nests along the loop, so that during the second survey all new nests could be recorded and their construction dated. Each time a fresh nest was identified, the ground immediately below it was searched for fresh (i.e. not entirely dry) chimpanzee feces. Opportunistic observations of nests away from these survey loops (and during initial visits to survey loops) were similarly recorded if leaves on observed nests were green, flexible, and not dry. Additional, opportunistic chimpanzee sightings were also recorded. For most analyses, nest and chimpanzee observations were lumped together, even though these two types of observation do not indicate precisely the same information about chimpanzee behavioral ecology (Strier 2003). Nests indicate where chimpanzees sleep overnight, while diurnal observations indicate where chimpanzees are active during the day, whether feeding, traveling, resting, or otherwise. Since the present research is meant to assess generalized chimpanzee distributionand not specifically the distribution of nesting, feeding, or other behaviorsin relation to various environmental features, lumping nest and chimpanzee observations together serves to increase sample size. This is important because the number of observations was low during some months and in some parts of the research area. Fecal contents. To estimate chimpanzee diet, fecal samples were collected during surveys (and opportunistically when found fresh and associated with a fresh nest) and dissected.

239 Following Moreno-Black (1978), McGrew et al. (1988), and Tutin and Fernandez (1993), the steps in this data collection effort were: 1) Chimpanzee feces identified based on: a) direct observation of defecation, or b) size, shape, texture, and location, being found fresh below a fresh nest; 2) Contaminating plant debris and insects removed from feces; each sample sealed individually and stored in plastic bag until dissection; 3) Fecal samples placed individually in 1-mm mesh bag and agitated in water to dissolve fecal matrix; 4) Items <1 mm diameter (e.g. Ficus seeds) floating in water skimmed out using cloth strainer (pore size <<1 mm); 5) Fecal material remaining after dissolution placed on metal plate and sorted; 6) Each item in this material identified through: a) prior familiarity, b) comparison with fresh plant specimens, c) comparison with items previously collected in fecal samples, or d) identification by taxonomist (for termites only); 7) Abundance of each item per fecal sample based on visual estimation of its volume after dissolution (cf. Basabose 2004), according to the following scale: 5 = >80% of sample, 4 = 50-80%, 3 = 20-50%, 2 = 1-20%, 1 = <1% (i.e. negligible volume); and 8) Samples of each item collected, dried, and stored for comparison with subsequent samples. Additionally, direct observation of chimpanzees or feeding remains, as described above, indicated consumption of other food items. Vegetation composition. Vegetation sampling used in this research is described fully in Chapter 4. To summarize, vegetation sampling occurred at 217 sites with disturbance histories

240 determined primarily through interviews of local residents. At each site, 0.1 ha was sampled according to Gentrys methodology (ten 2 (50 m plots: Phillips & Miller 2002). In these plots, all woody plants 2.5 cm diameter at breast height (DBH) were identified to species, measured (DBH only), and tallied. These tallies served as the bases of a hierarchical clustering analysis that identified the fifteen vegetation types listed in Table 1 (p. 264; see also Chapter 4). Relative dominance (basal area per species divided by basal area for all species), proportional abundance (number of individuals per species divided by total number of individuals), and relative frequency (number of sites in which a species was present divided by total number of sites) were calculated for each species per vegetation type. An importance value (IV) was calculated per species per vegetation type by adding its relative dominance, proportional abundance, and relative frequency scores. Plant phenology. The amount of fruits, flowers, and leaves borne by each individual plant observed during vegetation samples was assessed separately, using the following scale: 0=no fruits, flowers, or leaves; 1=less than 10 ripe fruits or fresh flowers, or less than 25% of the canopy with fresh leaves; 2=10-50 ripe fruits or fresh flowers, or 25-75% of the canopy with fresh leaves; 3=greater than 50 ripe fruits or fresh flowers, or greater than 75% of the canopy with fresh leaves. Distribution of biophysical features. Mapping permanent surface water sources and vegetation patches with high abundance of chimpanzee food plants was meant to estimate the spatial structure of biophysical features presumed important to chimpanzees. The distribution of permanent surface water was determined through interviews of local residents and foot surveys. Nine men, resident Maninka farmers and hunters, were interviewed to identify all locations where surface water is permanently available in the research area.

241 Permanent sources were considered those that have never been dry in living memory. The location of each permanent source was determined with a GPS unit, and the following characteristics recorded: type of source (i.e. spring, seep, or depression in drainage channel), abundance of water, vegetation cover, and human use. Foot surveys during the 2004 dry season (January-June) along all drainage channels and edges of all outcrops did not locate any additional, permanent water sources. Second, vegetation patches with a high abundance of chimpanzee food plants were mapped based on vegetation samples (described above) and a census of baobab trees (Chapter 5). Specifically, the following sites were mapped as points corresponding to each sites approximate center: 1) all patches of forest vegetation (i.e. vegetation types 1 and 2: Table 1, p. 264) larger than c.1 ha; 2) all sites that have not been occupied or farmed by humans since 1994 where 5 baobabs 100 cm diameter at breast height occur in an area 100 m 100 m; and 3) any other sites where woody plants with fruits eaten by chimpanzees (as identified through fecal samples or direct observation) were densely gregarious in an area 100 m 100 m. The disturbance history of all sites was determined through interviews of local residents (see Chapter 2). These sites provide a highly simplified representation of the distribution of chimpanzee food-plant patches. This representation is probably grossly accurate, and highly conservative because most food-plant species are sparse outside of these sites (Table 3, p. 268; see also Chapter 4). Ethnographic interviews. The researcher interviewed indigenous residents of the research area to gain information on their knowledge of chimpanzee behavioral ecology. Interviewees (n=12) were all male hunters between the ages of c.30-70. Interviews were all informal, primarily conducted during normal conversations, in which the subject of wildlife behavior was a common subject. Notes on the content of conversations were written during or

242 immediately after conversations. To clarify information provided during interviews, multiple interviewees were independently asked similar questions about in order to gain multiple perspectives on specific topics and improve the accuracy of interview results (Werner & Schoepfle 1987). Results Diet composition and seasonality. Thirty-eight food items were identified in 71 fecal samples, including five items of animal origin (Table 2, p. 266). Observations of chimpanzees or feeding remains at sites where chimpanzees were observed revealed nine food items; for six of these items there was no identifiable fecal evidence. The mean number of items per fecal sample was 3.451.32, the range 1-6. Fourteen items were not identifiable to taxa, and the identification of three items is tentative. Most unidentified items were single seeds, but many fecal samples included unidentifiable vegetal fiber. Fecal samples were found during February-July and September-December, but no food item occurred in fecal samples in every month. Most food items occurred in samples during limited periods of time, with high abundance during these periods. For fruits, these periods corresponded with the fruiting seasons of relevant species (Figure 2, p. 259). The majority of food items identified are of plant origin (Table 2, p. 266). Fruit remainswhether seeds or identifiable fibersaccounted for 29 of 38 food items, including an unidentified type of grass seed and eight unidentified types of dicot seeds. Excluding undifferentiated fiber, two types of leavesa grass and a dicot, both unidentifiedwere present in samples, and fibrous remains of bamboo stems (tentative identification) were seasonally very abundant. Direct observation and feeding remains of bamboo stem consumption were also abundant. Two types of flower were found as well: Daniellia oliveri flowers are a seasonally

243 important dietary item, while another, unidentified flower (tentatively determined to Caesalpiniaceae) occurred in a single sample. Chimpanzees were observed to eat the cambium of three tree speciesprimarily Pterocarpus erinaceus, but also P. lucens, and Cordyla pinnataalthough identifiable fecal evidence for this consumption was absent. Notably, local residents recognize the cambium of C. pinnata as a famine food (K. Dembl, personal communication, 3 Dec 2004). Of the five items of animal origin found in fecal samples, only twotermites of the genus Macrotermes and bee larvae (Apis africana, tentative identification)can be positively identified as dietary components. Two of the other threea tentatively identified roundworm (Ascarias spp.) and a mass of small, unidentified wormsare apparently parasites, while hairs observed in one sample may have been from a chimpanzee, although these were light in color. Four distinct periods can be identified over the course of the year in terms of chimpanzee diet (Table 2, p. 266). First, in February-March, most fecal samples were composed primarily of one of six food items, three of which are not fruits. This period had the lowest availability of ripe fruits based on phenological observations (Figure 2, p. 259). The mean number of items per fecal sample for February-March was 3.061.11. Second, in April-July, fruit remains dominated most fecal samples; this period also had fairly high availability of fruits, although few of the fruits that were abundant in fecal specimens were available (Figure 2, p. 259). The mean number of items per fecal sample was 4.081.50. The number of samples collected during this period the hot, dry seasonwas relatively low, because dry environmental conditions meant that many observed fecal remains were dry, and thus did not meet the fresh standard for collection. No data was collected in August. Third, in September-October, diet composition is almost entirely different from that in January-July. Most fecal samples were composed primarily of either

244 bamboo stem fiber, or remains of the fruits of Spondias mombin, Grewia bicolor, or Cissus populnea. These fruits were highly abundant during September-October (Figure 2, p. 259). The mean number of items per sample in September-October was 3.041.14. Finally, diet composition in November-December had composition similar to that of both September-October and February-March. The availability of fruit foods also showed some overlap between these periods (Figure 2, p. 259). The mean number of items per sample was highest in NovemberDecember (4.211.31). Diet composition certainly varies somewhat from year to year depending on the abundance of food sources. Local residents considered 2004 a notably poor year for edible wild fruit productivity generally, and especially for baobab and Hexalobus monopetalus. No comparative data is available to evaluate this impression. High percentages of the number of baobabs observed in September-December had ripe fruits. However, H. monopetaluswhich has sweet, moist fruitoccurred in only one fecal sample, and of 232 H. monopetalus individuals observed during vegetation sampling, only one (with <10 ripe fruits, on 18 Sep 2004) was observed with fruits. Observed diet composition (Table 2, p. 266) may be more indicative of chimpanzee behavior in years when the abundance of fruits is relatively low. Distribution of water sources and food patches. Permanent surface water sources and chimpanzee food patches are both most abundant along rock outcrops (Figure 2, p. 259). Water sources along cliffs are all springs or seeps coming from exposed sedimentary layers in sandstone bedrock. Thus, in the southern portion of the research area there are no permanent water sources along outcrops, because these are dolomite, whose hydrogeology differs from sandstone (DCTD 1990). No current human use of water sources along cliffs was observed or indicated in interviews. However, interviewees stated that past residents of several abandoned

245 settlements heavily used several of these sources prior to about 1940. In the relatively flat areas away from outcrops, nearly all water sources are depressions in seasonal streambeds. Humans regularly use most of these waterholes, primarily to provide water to domestic livestock. Most forest patches (vegetation types 1 and 2) occur along rock outcrops, although two occur in steep, narrow ravines incised into the edge of ferricrete hardpans, in the southern portion of the research area (Figure 2, p. 259). Most chimpanzee food patches identified more than c.100 m from a rock outcrop were baobab groves associated with abandoned settlement sites, although only a small proportion of abandoned settlement sites host a large number of large baobab individuals. Most sites have either a small number of large baobabs (2), or have only small individuals. Finally, particularly in the southern portion of the research area, some food patches occur along seasonal streams where food plants, especially Ficus spp., occur gregariously in bamboo thicket vegetation. All vegetation types include species for which evidence was collected of chimpanzee consumption, but chimpanzee food plants have highest importance values in gallery forests (vegetation types 1 and 2: Table 3, p. 268). If importance values of all food plants are used to estimate the abundance of food plants in vegetation types, several types of woodland and bamboo thicket are nearly as important sources of food plants as gallery forest, primarily because of the high relative frequency and proportional abundance of Pterocarpus erinaceus, and the high basal dominance of bamboo in many vegetation types. However, if only plants with fruits eaten by chimpanzees are retained in this analysis, food plants are clearly most abundant in gallery forests (Table 3, p. 268). Notably, the vegetation types with the next highest importance values for fruit-food plants are: 1) a type of woodland vegetation that occurs exclusively at abandoned settlement sites, and 2) bamboo thicket vegetation.

246 Distribution of nests and chimpanzee observations. A total of 695 nests in 224 nest groups and 131 chimpanzees in 48 groups were observed throughout the research area (Figure 3, p. 261). At this scale, chimpanzees and nests were observed in two broad patchesthe northern and the southern parts of the research areathat are separated by a gap several kilometers wide. This gap corresponds to an area of relatively flat topography without bedrock outcrops. As noted above, there are topographical and geological differences between the northern and southern portions of the research area. The median number of nests per group differs between the northern and southern areas, for all months and for three of four multi-month periods (Table 4, p. 269). A two-tailed MannWhitney U test (Bailey 1995) shows that the difference in nest group size is significant for the all-months medians (z=3.02, p<0.01). Difference in nest-group size between these two areas was also significant (z=2.53, p<0.05) for the September-October period. Within the broad northern and southern patches, nest groups are distributed patchily in dense clusters at a limited number of sites where chimpanzees build nests repeatedly throughout the year (Figures 3 & 4, pp. 261 & 262). Many of these sites occur along cliffs, but others occur some distance away from cliffs. The majority of nest groups were observed in vegetation types characteristic of cliff habitats, although the proportion of nests groups in cliff habitats was lower in the southern area than in north (Table 5, p. 270). Most other nest groups were observed in woodland habitats, although in the southern area a substantial minority occurred in bamboo thicket vegetation. The proportion of nests occurring in different habitats changed during the year. The occupation of some nest group sites in woodland areas was strongly seasonal, whereas others were occupied throughout the year (Figure 4). Most nest group sites along cliffs were occupied during all

247 seasons. Notably, most nest groups observed in woodland vegetation were in abandoned human settlement sites (Table 5, p. 270). Indeed, baobab groves at abandoned settlement sites are the most abundant type of food patch away from cliffs (Figure 3, p. 261). Finally, most nest groups were closer to food-plant patches than to permanent surface water sources (Table 6, p. 271; Figures 3 & 4, pp. 261 & 262). The distances between nest groups and either food-plant patches or permanent water sources varied somewhat from season to season. Notably, during the period April-Julythe hottest, driest period of the yeara greater proportion of nest groups were nearer to permanent water sources than during any other period. The greatest proportion of nest groups nearer to food patches occurred in the period November-December. Discussion Chimpanzee geography. The research area appears to include parts of the home ranges of two separate chimpanzee groups. The strongest evidence for this is the difference in nest group size between the northern and southern portions of the research area for all months of observation (Table 4, p. 269), which likely indicates different chimpanzee social organization in the two areas (Baldwin et al. 1981). While environmental differences between the northern and southern portions may contribute to different nesting behavior (cf. Baldwin et al. 1981)in particular, large predators, especially lions, are more abundant in the southern portion (Duvall, unpublished data from 2004; Duvall & Niagat 1997)it is unlikely that chimpanzees belonging to a single group would display such strong behavioral variation in moving from one portion of the landscape to another (McGrew et al. 1996). Local residents also believe that different groups occupy these areas, and report that it is very rare to see chimpanzees in the gap between the two areas.

248 Difference in nest group size between the two areas during the four multi-month periods displayed no or low significance (Table 4, p. 269). This is probably because the number of observations per period in the southern area was insufficient to gain an accurate estimate of median nest group size. Habitat differences between nest groups in the northern and southern areas (Table 5, p. 270) probably reflect environmental differences between these areas, rather than behavioral differences between chimpanzee groups. Sandstone outcrops dominate the northern portion of the research area, while the southern portion is relatively flat with just two small dolomite outcrops. This geology leads to topographical and, more importantly, hydrogeological differences between the northern and southern areas. These differences directly affect chimpanzees through the distribution and abundance of permanent water sources, and indirectly through the distribution and abundance of vegetation types. There are fewer, more widely dispersed permanent water sources in the south, and most of these are depressions in seasonal streambeds, occurring in relatively flat areas away from rock outcrops. Chimpanzee food patches are more widely dispersed as well, including in bamboo thicket vegetation along seasonal streamsan uncommon location for food patches in the northern part of the research area. Chimpanzees appear to select nesting sites based on the location of food patches because most nest groups, as well as observed chimpanzees, were nearer to food patches than water sources (Table 6, p. 271). However, many food patches and water sources occur near one another along rock outcrops, which means that the cost, in terms of increased distance to water, of nesting in a cliff-side food patch may be minimal. Indeed, there appear to be more permanent water sources in the research area than assumed by previous researchers (see Moore 1986; see

249 Moore 1985), meaning that water availability may not constrain chimpanzee ranging behavior as much as presumed, at least in areas with sandstone outcrops. Elsewhere, such as in the southern portion of the research area where there are not sandstone outcrops, water availability may affect chimpanzee distribution more significantly. Perched aquifersthe water held in sandstone outcropsis directly important to chimpanzees as the source of most permanent surface water in the research area, and indirectly important as a reason why forest patches occur along sandstone cliffs (see Chapter 4). The Manding Plateau area of southwestern Mali is important biogeographically as an area where numerous rare and relict plants occur, particularly associated with sandstone outcrops (Jaeger 1959). Cliffs in general, and sandstone cliffs in particular, represent ecologically very stable habitat where conditions change relatively little over millennia (cf. Larson et al. 2000). As a result, sandstone outcrops host numerous rare and relict plant and animal populations in arid to semi-arid areas around the world (Bowman et al. 1990; Bowman et al. 1988; Danin 1999; Jaeger & Winkoun 1962; Woodford 2000), and in southwestern Mali in particular (Duvall 2001; Jaeger 1956; Jaeger 1959; Jaeger 1966; Jaeger & Jarovoy 1952). Some chimpanzees in southwestern Mali are strongly linked to sandstone outcrops, and should be considered a component of the rare and biogeographically distinctive biota of the Manding Plateau. The abundance of water sources and food patches are both reasons why cliffs are the habitat chimpanzees most frequently use. First, permanent water sources are concentrated along sandstone outcrops, which links chimpanzees to these outcrops especially during the dry season. Permanent surface water away from outcrops occurs primarily in deep depressions along drainage channels, and humans have occupied or used most of these sites for decades to centuries. The availability of groundwater to trees adapted to growing in cracks in

250 sandstone outcrops also helps create the conditions necessary for cliff-side forests (see Chapter 4). Microclimate in these forests is cool and humid throughout the year, even in the dry season (Jaeger 1956). Indeed, chimpanzee and nest observations were most common cliff-side forests during the dry season, January-July (Table 5, p. 270). Second, many plants associated with sandstone outcrops are important food sources for chimpanzees, especially during September-October. Of course, with the present data it is not possible to say whether chimpanzees frequently use cliff habitats because of the abundance of food plants these habitats host, or if chimpanzee food plants are abundant in cliff habitats due to seed dispersal by chimpanzees. Nonetheless, several food plants are strongly abundant in cliff habitats and rare elsewhere in the landscape. In particular, fruits of the shrub Grewia bicolor are an important food source (Table 2). This plant dominates the edges of forest patches on sandstone outcrops (Duvall 2001; Jaeger 1950a; Jaeger 1956), and is strongly associated with bare sandstone outcrops (Table 3, p. 268; see also Chapter 4). The fruit-food plants Cissus populnea, Cola cordifolia, Cordia myxa, Diospyros mespiliformis, Erythrophleum suaveolens, Saba senegalensis, Sarcocephalus latifolius, Sorindeia juglandifolia, and Spondias mombin are also most abundant in cliff-side gallery forests, but are less restricted to sandstone outcrops (Table 3, p. 268; see also Chapter 4). Since cliffs provide a wide range of microhabitat conditions ranging from xeric to mesic (Larson et al. 2000), cliff habitats host the greatest variety of food plants, including many species that are characteristic of woodland and bushland, and not just forest, vegetation. Foods from plants characteristic of cliff habitatsespecially G. bicolor and S. mombinare particularly important in the period September-October (Table 2, p. 266). Chimpanzees may prefer these foods, or the proximity of most cliff-side food patches to permanent water sources, because the highest proportion of baobabs with ripe fruit occurred in

251 September-October, at the same time as peaks in fruit availability for G. bicolor and S. mombin (Figure 2, p. 259), but evidence for baobab fruit consumption did not appear until after G. bicolor and S. mombin were no longer fruiting (Table 2, p. 266). While cliff habitats are ecologically stable (Larson et al. 2000), perched aquifers in sandstone outcrops ultimately depend upon recharge from rainfall (Dunne 1990). Africa is the least-studied, populated continent with regard to the potential effects of climate change (IPCC 1996). While minor increases in average annual precipitation are predicted for semi-arid West Africa (Hly et al. 2006), the timing of rainfall is expected to change more substantially (Caminade et al. 2006). The effects of such change are not known precisely, but forest vegetation is expected to be most sensitive to changes in rainfall amount and timing (Hly et al. 2006). Additionally, changes in precipitation regimes will substantially change runoff rates, and thus rates of groundwater recharge (de Wit & Stankiewicz 2006). Considering that West Africa has experienced increased aridity in the past 160 years (Schngart et al. 2006), forests reliant upon groundwater may already face significant climate-change induced stress. Monitoring groundwater flow rates in sandstone outcrops and the composition, structure, and distribution of forests on these outcrops should be considered an important aspect of chimpanzee conservation and management efforts in Mali. Chimpanzees and humans. Humans and chimpanzees have shared Sudanian West Africa for thousands of years, and the results show that chimpanzees are ecologically adapted to some types of agricultural landscape. African farmers have profoundly altered plant distributions through migration, settlement, and cultivation (O'Brien & Peters 1998). In southwestern Mali, human settlement has led to the development of baobab groves at settlement sites (see Chapter 5), and to the introduction of other edible fruit-bearing, African trees that were originally absent

252 (Maranz & Wiesman 2003). These baobab groves are important to the trees reproduction because large (i.e. reproductive) individuals are most abundant, and significantly overabundant relative to other habitat types, in abandoned settlements and fields (see Chapter 5). Other wild fruit trees are present at abandoned settlement sites, due in part to human activities, but also because the environmental conditions at these sites are suitable for trees with seeds dispersed by animals that feed on the fruits of trees, like baobab, that survive settlement occupation (see Chapter 4). Chimpanzees are amongst those animals that benefit from the density and abundance of fruit trees at some abandoned settlement sites, and contribute to the further development of these fruit-tree patches by dispersing seeds to these sites. Chimpanzee use of abandoned settlements is strongly seasonal, corresponding to the abundance of food sources in different habitat types (cf. Naughton-Treves et al. 1998). Half (50.8%) of all nest groups observed during November-December were in abandoned settlement sites, while during other portions of the year, only 6.0-15.9% were in such sites (Table 5, p. 270). Chimpanzee use of abandoned settlements corresponds to the period when baobab fruit comprises a significant portion of the animals diet (Table 2); baobabs are most abundant in abandoned settlements (Table 3, p. 268; see also Chapter 5). During other parts of the year, most important food items come from plants associated with cliff habitats (Tables 2 & 3, pp. 266 & 268). Notably, though, chimpanzees do not appear to increase their use of baobab groves at abandoned settlements in direct correspondence to increases in the availability of ripe baobab fruit. The highest proportion of baobabs with ripe fruit occurred in September-October (Figure 2, p. 259), but sharply increased use of abandoned settlement sites did not occur until NovemberDecember, after two important food plants associated with cliff habitatsGrewia bicolor and Spondias mombinwere no longer fruiting (Figure 2, p. 259). Similarly, evidence for

253 chimpanzee feeding on figs and termitesboth of which represent fallback foods for chimpanzees, and frugivores more generally, when fruit abundance is low (McGrew et al. 1988; Terborgh 1986)increased only after G. bicolor and S. mombin had finished fruiting (Table 2, p. 266), even though figs (and, presumably, termites) were available throughout the year (Figure 2, p. 259). Baobab groves at abandoned settlement sites may represent a fallback source when fruit food is not abundant in cliff habitats. Past observations of chimpanzee ecology in Mali further support these results. Most published observations have come, coincidentally, from the period November-January, when observers found high abundance of chimpanzee nests near large baobabs (Granier & Martinez 2004; Moore 1985). In contrast, Pavy (1993), who surveyed the area in February-April 1992, made no mention of chimpanzee or nest observations near baobabs and had few observations in woodland areas more generally; he did find a strong correlation between nest distribution and topography. [T]hree times out of four he found nests when encountering a cliff or steep slope along line transects (Pavy 1993: 19). While he did not describe vegetation, his data on the identification of trees hosting nests (J.-M. Pavy, personal communication, December 2002) suggest that at least 60% of nests were observed in vegetation comparable to vegetation types 1 or 2 (Table 1, p. 264). Local residents also recognize that chimpanzees use abandoned settlements primarily during baobab fruiting season, which is considered an unintentional outcome of settlement practice. This knowledge has gained importance because of the influx of chimpanzee researchers who have visited Solo since 1983. When the primatologist James Moore visited Mali in 1983 and asked the village of Solo to provide a guide for his search for chimpanzees, Solos leaders agreed because Moores proposition seemed to hold promise for future benefits

254 for the village (F. Dembl, personal communications, March 2004). After discussion amongst themselves separate from conversations held with Moore and his Malian colleague, Solos hunters decided that the guide should lead him to several baobab groves at abandoned settlements sites since baobabs were fruiting at the time (F. Dembl, personal communications, March 2004). The guide did not tell Moore that these baobab groves were abandoned settlement sites (F. Dembl, personal communications, March 2004). The hunter recalls finding fresh nests at most of these groves, and Moore (1985: 60) reported that many nests were observed near fruiting baobabs. While the group also visited other locations, including a bedrock outcrop south of Solo, the decision of Solos hunters to focus search efforts on abandoned settlement sites was based on their knowledge of the seasonality of human-baobab-chimpanzee interaction. Similarly, the primatologist Pascal Gagneux visited Solo in November 1995, seeking chimpanzee nests from which hair samples could be collected (cf. Gagneux et al. 1999). Since Gagneuxs visit was several weeks earlier in the year, his guide decided to visit primarily forest patches along the sandstone outcrop north of Solo, but also visited two baobab groves at abandoned settlement sites (K. Dembl, personal communications, November 2004). They found chimpanzees in both types of site, but more abundantly in cliff-side forests (K. Dembl, personal communications, November 2004). Maninka farmers do not aim to attract chimpanzees to settlement sites, because chimpanzees are not considered a valuable resource, as a source of meat or other products (Duvall & Smith 2005; Granier & Martinez 2004). However, occupied and abandoned settlement and field sites are often managed with an explicit goal of attracting certain frugivorous or omnivorous game animals (cf. Fairhead & Leach 1996), such as duikers (Cephalophinae), bushbuck (Tragelaphus scriptus), and warthog (Phacochoerus aethiopicus). For instance, fig

255 trees (Ficus spp.) are often maintained in fields and around occupied and abandoned settlement sites because fallen figs attract these animals; humans generally do not eat wild figs. Few wild animals consume baobab fruits because the shells are too difficult to break open; baobabs are maintained because of their utility to humans, in addition to their historical and spiritual significance (see Chapter 5). Chimpanzees are able to break open baobab fruits (Marchant & McGrew 2005), and their use of abandoned settlements is an unintended, but clearly understood, outcome of Maninka settlement practice. Human and chimpanzee use of baobab fruit represents potential competition between the species. Residents of Solo do not consider chimpanzees a pest, though, because humans tend to rely on baobabs in occupied settlements and fields, and abandoned settlements near primary footpaths. Chimpanzees visit only abandoned settlements, and generally only those distant from occupied settlements and primary footpaths (Figure 3, p. 261). An aggressive response by conservationists to potential human-chimpanzee competition for baobab fruit would perhaps entail prohibition of all baobab fruit collection outside of occupied settlements and fields. This response would overlook the importance of wild fruit to rural livelihoods, and hinder the cause of chimpanzee conservation by alienating rural people who have great knowledge of chimpanzee behavior and the ability to either protect or extirpate Malis chimpanzees. Knowledgeable local residentsindigenous huntersrecognize that humans and chimpanzees share wild fruit resources by foraging in different parts of the landscape. Most local residents would accept policies that clearly specify open and closed areas for wild fruit collection if these are developed, explained, and enacted in a way that reflects indigenous knowledge, and not just the global rarity of chimpanzees and the conservation significance of Malis population. Conclusion

256 In southwestern Mali, chimpanzees frequently use abandoned settlement sites during the time of year when baobab fruit is a significant component of their diet. Baobab groves at settlement sites develop directly and indirectly as the result of human activities, which means that humans have effectively increased the distribution of chimpanzee food plants across the landscape, creating seasonally important habitat patches. Baobab is less abundant elsewhere in chimpanzee range, so it is unclear how widely chimpanzees and humans interact in this way because. However, similar relationships between humans, plant distributions, and wildlife probably occur widely in Africa where human settlement practice and history have led to the development of distinct vegetation patches at abandoned settlements. Indirect effects of human activities on wildlife certainly have a deep evolutionary history in Africa, and conservationists should recognize these relationships in order to develop policies that more accurately reflect land-use ecology, and minimize potential, negative, long-term effects of the forced alteration of customary land-use practices. While short-term threats to chimpanzee survival must remain a priority, the biological and, in some aspects, ecological similarity between chimpanzees and humans may mean that human activities can serve as an asset to the long-term viability of chimpanzee populations and their habitat. Wildlife managers must recognize this possibility, and carefully evaluate how human selection and use of wild fruit trees may, over longer timescales, prove beneficial to chimpanzees.

257 Figures and tables for Chapter Five

258 Figure 1. Western Mali, showing chimpanzee distribution and research area. Inset map of Africa shows area represented by main map. Chimpanzee distribution from Duvall et al. (2003).

259 Figure 2. Phenology of chimpanzee fruit-food plants. Numbers to the left of graphs show the percent of all individuals observed per month with any ripe fruit; numbers below graphs show the number of individuals observed with any ripe fruit over the total number of individuals observed per species. Individual graphs are shown only for species with fruits eaten by chimpanzees that composed >20% of >1 fecal samples. The graph Ficus spp. Fig lumps observations for 13 fig species, including F. cordata, the only species for which there is speciesspecific evidence for its consumption. The graph All other confirmed fruit foods lumps observations for the 14 species with fruits eaten by chimpanzees that occurred in at least one fecal sample (i.e. Diospyros mespiliformis, Saba senegalensis, Cordia myxa, Cola cordifolia, Lannea spp., Sorindeia juglandifolia, Hexalobus monopetalus, Parkia biglobosa, Vitex madiensis, Tamarindus indica, and Boscia angustifolia), or for which there was direct evidence for chimpanzee consumption (i.e. Erythrophleum suaveolens). The item Lannea spp. includes observations of L. acida, L. microcarpa, and L. velutina. The graph Candidate fruit foods lumps observations for 23 other species encountered in vegetation samples that either have fruits considered edible by local residents, or are congeneric with plants having fruit remains occurring in fecal samples (i.e. Annona senegalensis, Borassus aethiopum, Cordyla pinnata, Detarium microcarpum, Diospyros abyssinica, Gardenia erubescens, Grewia flavescens, G. lasiodiscus, G. venusta, Landolphia heudelotii, Malacantha alnifolia, Manilkara multinervis, Pachystela brevipes, Parinari curatellifolia, Raphia sudanica, Sclerocarya birrea, Strychnos spinosa, Trichilia emetica, Vitellaria paradoxa, Vitex doniana, Ximenia americana, Ziziphus mauritiana, and Z. mucronata).


261 Figure 3. Biophysical features in the research area, and observations of chimpanzees and nests.
a) Biophysical features and survey loops
1 4


5 3

Seasonal streams Bafing Reservoir Survey loops 6 Clif Occupied settlements Permanent surface water sources Food-plant patches: cliff forests, baobab groves, other (primarily bamboo thickets) Primary footpaths Chimpanzee sightings Chimpanzee nest groups 0 5 km

b) Chimpanzee and nest observations

262 Figure 4. Seasonal observations of chimpanzees and nests.



Table 1. Floristic vegetation types present in the research area. For full description of these vegetation types, as well as explanations of the vegetation type codes and names used below, see Chapter 4. These vegetation types are based on floristic composition, but most have strong environmental correlates, indicated in the Habitat description column. Vegetation structural categories (i.e. forest, thicket, woodland, and wooded grassland) reflect Lawessons (1995) definitions.

265 Code Name 1 Cola cordifolia-Spondias mombin forest 2 3 4 5 6 7 8 9 10 11 12 13 14 15 Gilletiodendron glandulosum-Hippocratea indica forest Oxytenanthera abyssinica (bamboo) thicket Xeroderris stuhlmanniiBombax costatum wooded grassland Ferricrete wooded grassland Combretum nigricans wooded grassland Crossopteryx febrifuga wooded grassland Terminalia macroptera woodland Rupicolous bushland Pterocarpus lucens-Gueira senegalensis bushland Dichrostachys cinerea bushland Acacia ataxacanthaCombretum micranthum woodland Terminalia macropteraVitellaria paradoxa woodland Pteleopsis suberosaHymenocardia acida woodland Pterocarpus erinaceusVitellaria paradoxa woodland Habitat description Topographically sheltered sites, in narrow ravines along rock outcrops. Most sites with permanent surface water and permanently wet soil. Topographically sheltered sites with extremely rocky (sandstone) soil. Many sites with seasonal surface water, all with seasonally dry soil. Strongly dominated by bamboo. Sites along seasonal streams that are not sheltered from fire. Steep, rocky slopes, in topographically high sites (i.e. away from drainage channels). Topographically high sites with gravelly, silty soil shallowly overlying a ferricrete horizon. Topographically high sites with shallow, infertile, and dry soil. Sites with silty soil that are seasonally waterlogged due to poor drainage. Topographically high sites with apparently low fertility. Topographically high sites with very rocky (sandstone) soil. Sites with naturally eroded, clayey loam soil, at seasonal drainage channel heads. Sites disturbed by past settlement. Vegetation dominated by species with edible fruits. These four woodland vegetation types share characteristic habitat: sites with deep, arable soil that is neither xeric nor highly mesic. Over 80% of these sites have been disturbed by past human settlement or cultivation.

266 Table 2. Seasonality of chimpanzee diet. Abbreviations: B=bark; Fl=flower; Fr=fruit; L=leaf; S=seed; St=stem; Unk.=unknown; vol.=volume. Food items are arranged from top to bottom in the temporal order in which evidence for their consumption was collected. Thus, items that were first encountered in fecal samples early in 2004 are found toward the top of the table, while items that were first encountered later in the year are found toward the bottom. Ficus cordata is not listed in this order, but is placed next to the entry for unidentified Ficus seeds. Tentative identifications of food items are given in brackets. Gray ovals indicate the volumetric composition of fecal samples (see key); black diamonds represent direct evidence of chimpanzee diet, through observations of chimpanzees. Each vertical column of gray ovals represents the contents of one fecal sample. Direct evidence is indicated in the column of the fecal sample whose collection date corresponds most closely with the date of the direct observation.


268 Table 3. Abundance of chimpanzee food plants in vegetation types. For Vegetation type codes, see Table 1 (p. 264). Species names have been abbreviated from those listed in Table 2 by retaining the first two letters of the genus and species names. Values shown are importance values (IVs) per species for each vegetation type. IVs <0.1 are represented by a tilde ~. Totals are the sums of IVs for all species per vegetation type. Fruits row sums IVs only for plants with fruits for which there is evidence of chimpanzee consumption (Table 2, p. 266).
Species Ad. di. Bo. an. Ci. po. Co. co. Co. my. Co. pi. Da. ol. Di. me. Er. su. Fi. spp. Gr. bi. He. mo. La. spp. Ox. ab. Pa. bi. Pi. th. Pt. er. Pt. lu. Sa. la. Sa. se. So. ju. Sp. mo. Ta. in. Vi. ma. Totals Fruits 1 0.1 0.4 0.4 1.2 0.4 0.1 1.5 0.4 1.3 0.7 0.1 0.5 1.0 0.1 0.1 0.2 0.1 0.7 0.5 0.2 1.2 0.1 11.3 9.9 2 0.1 0.7 0.3 0.4 0.1 3 4 0.4 0.1 0.2 5 6 0.4 0.1 Vegetation types 7 8 9

10 0.6

11 0.9 0.1 0.1 0.1 0.1 1.1

0.1 0.1 0.2 0.2 0.2 0.4 1.0 0.3 0.5 2.5 0.1 0.3 0.7 0.8 0.5 0.4

0.1 0.1 0.2 0.2

12 0.9 0.1 0.1 0.1 0.2 0.1 ~ 0.1 0.2 0.8 0.5 0.1 0.6 1.5

13 0.7

14 0.2 ~ ~

15 0.2 0.1

0.1 0.2 0.1 0.4 0.3 0.5 0.9 0.5 0.1 0.2 0.1 0.1 0.6 1.0 0.7 0.1 0.2 1.3 0.2 0.7 1.5 1.6 0.1 0.2 1.1 0.3 0.1 0.1 0.6 7.1 3.4 0.2 5.2 3.8 0.4 4.8 2.5 0.2 5.4 2.9 0.3 0.2 0.9 1.0 0.1 0.1 0.3 0.8 0.1 0.1 1.4 0.1 0.3 1.2 0.4 0.4 1.3 0.8 0.8 0.9 0.2 0.9 0.9 0.4 0.5 0.5 0.7 0.7 0.3 0.3 ~ ~ ~ 0.4 1.4 0.3 0.2 0.6 1.4 ~ 0.1 ~ 0.1 6.6 3.7 0.4 5.6 3.3 ~ 0.1 5.0 2.7 0.2 0.1 ~ ~ 0.3 1.1 0.3 0.1 0.8 1.7 ~ ~

0.6 0.2 1.2 1.1 0.3 0.8

0.2 0.1 1.0 0.3 0.3 1.1 1.1 0.1 0.1


0.9 1.4

0.6 1.6 0.1

0.6 0.8 0.2 0.1 0.9 0.9

0.8 0.9 0.1 1.0


0.1 0.2 0.1 0.1 6.4 5.2

0.2 0.1 ~ 0.1 5.7 3.4

8.6 7.8

8.3 4.9

5.2 2.9

3.5 1.4

6.1 2.5

269 Table 4. Summary statistics for nest group size. Abbreviations: n, number of nest groups observed; p, statistical significance. Mann-Whitney U test was used to compare median nestgroup size between the northern and southern portions of the research area. Significance shown only where p<0.05. All months n 224 median 2.5 n 198 median 2 n 26 median 4 p <0.01 Jan-Mar 50 3 44 3 6 4 Apr-Jul 63 2 60 2 3 2 Sep-Oct 37 2 32 2 5 4 <0.05 Nov-Dec 74 2 62 2 12 4

Entire area Northern portion Southern portion

270 Table 5. Nest group characteristics. Cliff habitats includes Vegetation types 1, 2, and 9 only (Table 1, p. 264). Woodland habitats include vegetation types 12, 11, 13, and 15 only (Table 1, p. 264). Observations made in Abandoned settlements are a subset of the observations made in woodland, and occurred 100 m from the center of an abandoned settlement site. Thicket habitat is vegetation type 3 only (Table 1, p. 264). Nests were not observed in other vegetation types. Area All months Entire Cliff 138 research (61.6%) area Woodland 79 (n=224) (35.3%) Abandoned 54 settlement (24.1%) Thicket 7 (3.1%) Northern Cliff 135 portion (68.2%) (n=198) Woodland 63 (31.8%) Abandoned 49 settlement (24.7%) Thicket 0 Southern Cliff 3 (11.5%) portion Woodland 16 (n=26) (61.5%) Abandoned 5 (19.2%) settlement Thicket 7 (26.9%) Habitat Jan-Mar 40 (80.0%) 7 (14.0%) 3 (6.0%) 3 (6.0%) 39 (88.6%) 5 (11.4%) 3 (6.8%) 0 1 (16.7%) 2 (33.3%) 0 3 (50.0%) Apr-Jul 48 (76.2%) 14 (22.2%) 10 (15.9%) 1 (1.6%) 46 (76.7%) 14 (23.3%) 10 (16.7%) 0 2 (66.7%) 0 0 1 (33.3%) Sep-Oct 19 (51.4%) 17 (45.9%) 9 (14.3%) 1 (2.7%) 19 (59.4%) 13 (40.6%) 9 (12.1%) 0 0 4 (100%) 0 0 Nov-Dec 31 (41.9%) 41 (55.4%) 32 (50.8%) 2 (2.7%) 31 (50.0%) 31 (50.0%) 27 (43.5%) 0 0 10 (83.3%) 5 (41.7%) 2 (16.7%)

271 Table 6. Distances between nest groups and water sources or food patches. All months 224 48 44 (16.2%) 42 (15.4%) 186 (68.4%) 216 578 Jan.-Mar. Apr.-Jul. 50 7 8 (14.0%) 12 (12.1%) 37 (64.9%) 171 476 63 18 18 (22.2%) 11 (13.6%) 52 (64.2%) 255 619 Sep.-Oct. 37 17 5 (9.3%) 12 (22.2%) 37 (68.5%) 250 656 Nov.Dec. 74 6 10 (12.5%) 8 (10.0%) 62 (77.5%) 185 763

Nest group observations Chimpanzee observations Observations nearer to water source than food patch Observations 100 m to both water source and food patch Observations nearer to food patch than water source Median distance to nearest food patch (meters) Median distance to nearest water source (meters)

272 Chapter Seven: Conclusion For decades, researchers have recognized that humans have profoundly shaped African ecosystems by affecting both the abundance and distribution of plant species. Earlier analyses of human impacts on African ecosystems tended to underscore negative impactssuch as deforestation or other declines in plant density and/or diversitywhile more recent works have tended to draw attention to positive impactsgenerally increases in (or the stability of) plant density or diversity. In both cases, past studies have often overgeneralized the spatial extent of anthropogenic impacts, either across focal landscapes or more broadly across the continent. In short, Africaand West Africa in particularhas often been represented as a place where anthropogenic disturbance is the primary determinant of ecosystem composition and structure. In contrast, this dissertation shows that human activities are not necessarily the dominant factor determining these ecosystem characteristics in rural landscapes. Humans act within biophysical and sociocultural contexts that create variability in disturbance intensity, and that determine the significance of human disturbance as a source of ecological heterogeneity and change. Biogeographers recognize that similar vegetation does not necessarily develop in ecologically similar sites (McCune & Allen 1985). Likewise, human-environment scholars must recognize that the intensity and significance of ecosystem response to human disturbance depends on the specific context in which disturbance occurs. Thus, human environmental impacts varyacross landscapes as well as between different landscapesdepending on the biophysical and sociocultural contexts in which disturbance occurs. Many generalized relationships between human activities and biodiversity resources are accurate in many landscapes, but some of these generalized relationships may be inaccurate in portions, or all, of some landscapes.

273 The importance of recognizing local context to understand human environmental impacts is the core of many geographical critiques of conservation practice. This dissertation contributes to this widely shared geographical argument. More importantly, however, it supports two conclusions on how human-environment knowledge is created and applied. First, this dissertation shows that the dominance of certain scientific traditions over others may create an inaccurate appearance of certainty in scientific assessments of human-environment relationships. Second, this dissertation shows that the spatially fixed and temporally absolute boundaries generally used in conservation practice are inadequate for managing human-environment interactions in strongly heterogeneous environments. These conclusions are discussed separately in the following two sections. Scientific knowledge creation and transmittal. Why have past researchers failed to recognize adequately the importance of context in analyses of anthropogenic environmental change in West Africa? This dissertation contributes to recent work on the role scientific discourse has played in the development and maintenance of dominant, but inaccurate, environmental narratives of human-environment relationships in Africa (e.g. Bassett & Crummey 2003; Cline-Cole & Madge 2000; Fairhead & Leach 1998; Leach & Mearns 1996). Many of these challenges to dominant environmental narratives have shown that indigenous knowledge can offer more accurate representations of biophysical reality than environmental sciences have in the West African context (Bassett & Koli Bi 2000; Fairhead & Leach 1996; Richards 1985). Indeed, Chapter 3 of this dissertation shows that Maninka categorization of land cover, for instance, is considerably more detailed than that of natural resource scientists. Nonetheless, counterpoising scientific and indigenous knowledge oversimplifies the range of viewpoints people have on environmental questions (Agrawal 1995; Atran 1990). In particular,

274 political ecologists have often overlooked divergent traditions that exist within the broad scientific community (Turner 2005). Science has been portrayed monolithically in many recent challenges to dominant environmental narratives in Africa, although some have acknowledged differences between scientists concerned with particular problems (Bassett & Boutrais 2000; Beinart 1996; Duvall 2003; Richards 1985). In contrast, this dissertation shows that science does not necessarily offer a single, dominant narrative, against which only local knowledge may be offered as counternarratives. Science may include neglected literatures and viewpoints that can effectively counter dominant narratives. Many of the findings of this dissertation build upon earlier academic work that has been neglected in the Malian, if not African, research literature. Two key examples may be drawn from the findings. First, conservationists in southwestern Mali have focused their efforts on rare trees associated with sandstone bedrock, and on chimpanzees associated with these trees. To explain why these trees are associated with sandstone outcrops, conservationists have relied on the anthropogenic deforestation theory of African vegetation history, which has dominated natural scientific discourse in Africa for decades (Chapter 4). This theory depicts the patchy distribution of the endemic tree Gilletiodendron glandulosum and the forest type it dominates as evidence for past deforestation caused by indigenous land management practices (Duvall 2003). However, anthropogenic deforestation theory fails to incorporate abundant findings from West Africa (e.g. Chudeau 1917; Larminat 1927) and worldwide (Larson et al., 2000) that show clearly how cliff habitats create ecologically very stable microhabitats, and are likely to host rare species (Chapter 4). Second, Chapter 1 shows that representations of Maninka settlement in the Bafing area as an expanding frontier neglects not only local settlement history, but also a

275 literature on the wide occurrence of shifting settlement practice in Africa (de Schlippe 1956; e.g. Murdock 1967; Netting 1993; Richards 1978; Sidikou 1974; Stone 1996). The dominance of particular scientific viewpointssuch as anthropogenic deforestation theory, or frontier-style settlementon the geography of biodiversity has given the appearance of certainty in understanding how human activities pose threats to natural resources in southwestern Mali. Yet this dissertation, supported by neglected scientific literature, shows that biodiverse habitats do not necessarily represent the remains of pre-settlement landscapes humans have devastated through resource use. Instead, these habitats may reflect ecological conditions that exist independently of humans (Chapter 4), or they may represent the outcome of human activities (Chapters 5 and 6). Again, however, context matters. For instance, in the case of Polylepis forests in the Andessimilarly patchy habitats dominated by a narrowly endemic tree genusdiscursive inertia privileged biophysical conditions over human activities to account for the patchy distribution of these trees, a viewpoint that has recently been overturned by careful observations on Polylepis ecology and distribution (Fjelds 2002; Purcell et al. 2004). Science may be inherently political (Forsyth 2003), but it also claims to be scientific and thus can be evaluated on its own terms. Uncovering the scientific lineage of received ideas also exposes other scientific ideas, which compose alternate narratives about particular environmental questions. Human-environment scholars must recognize that much research of current interest and apparent novelty may have neglected antecedents, and must seek these antecedents in order to improve understanding of scientific knowledge transmission, as much as human-environment relationships. Boundaries and mobility in conservation practice. The most significant contribution this dissertation makes is to understanding the biogeography of human activities in the semi-arid

276 tropics, and how this affects the appropriateness of conservation practices in different landscapes. This contribution is grounded on recognition of three aspects of local context, underscored by this dissertation, that may lead to variability in the intensity or significance of human disturbance. First, different land useseven closely linked land uses, like settlement and cultivation in agrarian rural landscapesdiffer significantly as ecological disturbances because indigenous knowledge and practice varies according to land use. Specifically, vegetation analysis in Chapter 4 shows that settlement and cultivation have distinct effects on vegetation composition, because the goals of vegetation clearing for settlement differ from that practiced for cultivation. As argued in Chapter 2, settlement must be recognized as a distinct land use in natural resource conservation and management. Maninka farmers seek to eliminate most woody vegetation from settlement sites for the duration of site occupation, while in fields the goal is to reduce tree canopy cover, without killing trees, for the relatively short period of swidden use. These differences in land management practice favor different plant adaptations to disturbance: reproduction from seed is more important in settlement sites, while vegetative reproduction is more important in fields (Chapter 4). Variation in ecological knowledge and practiceas a function of land-use goals and as a characteristic of difference between individuals and societiescreates variation in the intensity of environmental impacts caused by humans. Second, biophysical factors that are significant in biotic variation at regional spatial scalessuch as soil and hydrology in West Africa and the semi-arid tropics more generally may limit the significance of human disturbance as a source of ecological heterogeneity in focal landscapes. Vegetation analysis in Chapter 4 shows that human activities affect vegetation composition, but also that edaphic characteristics are a more significant source of habitat

277 heterogeneity across the research area. Similarly, analysis of chimpanzee distribution in Chapter 6 shows that chimpanzees use anthropogenic habitat patches, but also that other, nonanthropogenic types of habitat are more frequently used throughout the year. In both cases, a spatial framework created by edaphic variation limits settlement and cultivation to specific portions of the focal landscape. Ecological heterogeneity in other parts of the landscape is not attributable to human activities. Furthermore, edaphic variation can limit the ecological significance of disturbance: the composition of disturbed vegetation may not differ significantly from other vegetation, especially if observed relative to the full range of plant communities present in the landscape. In other words, recognition of some anthropogenic disturbances either as types of event or as specific eventsmay be meaningful primarily as indications of human history and not of ecological heterogeneity, depending on which portions of a landscape and which ecological processes are of interest. Indeed, for Maninka farmers, manyang (fallows) are distinct because of their history and not necessarily because of vegetation, soil, or other biophysical characteristics (Chapter 3). Third, human influence on ecosystem structure and function must be understood in a long-term temporal context. Awareness of long-term timescales is particularly important in Africa because of its extremely long human history. Analysis of spatiotemporal relationships between humans, baobabs, and chimpanzees in Chapters 5 and 6 shows that the ecologies of these species are closely intertwined, both directly (for humans and baobabs, and for baobabs and chimpanzees) and indirectly (for humans and chimpanzees, in terms of habitat distribution, composition, and abundance). These linkages have arisen from a long history of interaction, and contribute to the emergent structure of the focal ecosystem. This history matters for the longterm, future ecosystem structure in the research area. As shown in Chapter 2, altering settlement

278 practice alters the spatial distribution of human impacts, and perhaps also the intensity of impacts in affected areas. Spatial change in settlement practice may directly impact biodiversity resourcesif, for instance, human use of biodiverse habitats increases in areas where the number of people has increased because other areas are closed to settlement (Chapter 2)but indirect effects may be more significant in the long termif reduced frequency of settlement establishment and abandonment reduces recruitment for wild fruit trees across the landscape (Chapter 5). While the effects of specific disturbance events may fade over decades to centuries (Chapter 4), the anthropogenic disturbance regimethe cumulative effects of individual disturbance eventsover longer timescales may represent a mechanism that contributes to the maintenance of certain ecosystem characteristics. Recognizing how these aspects of local context may constraint the spatial and ecological significance of human activities as environmental disturbances enables assessment of the appropriateness of spatial and temporal scales implicit in conservation practice. Of all conservation interventions, spatial and temporal scale is perhaps most strongly inherent in the establishment of protected-area boundaries. Protected-area boundaries are in virtually all cases spatially explicit, and generally also delineate absolute breaks in the spatial distribution of allowed human activities, regardless of temporal variation in human activities or biophysical conditions. However, these boundaries are created in landscapes where biophysical and sociocultural processes either are unbounded spatially, or occur within spaces that are not coterminous with protected areas. In either case, the rate and direction of environmental changes associated with biophysical and sociocultural processes are not necessarily consistent, so that the apparent appropriateness of a boundary may change over time. The inadequacy of rigid, absolute borders to encompass spaces that are ecologically meaningful either for humans or other

279 ecosystem components helps create conservation failures in landscapes where mobility is a crucial aspect of land use and livelihood security (Turner 2006). Human mobility has been most widely recognized in landscapes where pastoral livestock husbandry is practiced. In landscapes where shifting settlement and/or shifting cultivation are practiced, mobility is as important to land use and livelihoods as in landscapes where pastoral livestock husbandry is practiced, but the rate of mobility differs in settlement, cultivation, and pastoral husbandry practices (cf. Stone 1996). Simplistically, time periods over which mobility is evident in settlement practices are longer than those for cultivation, which are longer than those for pastoralismalthough there are longer-term dynamics in each type of land use that complicates such simple, generalized comparisons. In any case, conservation practices that emphasize rigid boundaries and absolute land-use delineations are poorly suited to managing spatial mobility and variation over time, whether these are associated with humans or other ecosystem components (Kozakiewicz 1995; Turner 2006). Clearly, short-term observations of human practices or biophysical conditionswhich have been central to the development of conservation theory and practice in West Africaare inadequate for understanding the spatial dynamism of land use and associated environmental changes. More importantly, the failure of conservationists to recognize long-term mobility in traditional land management has meant that conservation practitioners are poorly equipped conceptually to deal with environmental changes that develop over the course of decades, and not shorter periods. For instance, rigid boundaries are unable to adjust to slow or subtle shifts in the distribution of human activities or biodiversity resources, while absolute land-use delineations are unable to adapt to infrequent boundary crossings by humans or wildlife resulting from relatively short-lived environmental conditions regularly or widely separated in time, such

280 as seasonal change, or natural disasters. Conservationists must accept that: a) the importance of mobility increases for organismsincluding humansas landscape heterogeneity and patchiness increases (Kozakiewicz 1995); and b) accept that the distribution of human activities and biodiversity resources changes over timescales longer than it is feasible to observe before making conservation decisions. In other words, rigid and absolute boundaries are ecologically alien to heterogeneous landscapes, and the introduction of such boundaries will sooner or later disrupt natural and human ecological processes, many of which scientific observers may be unaware because of their long periodicity. Thus, in markedly heterogeneous landscapessuch as semi-arid tropical woodlands and grasslands, but not necessarily tropical rainforests conservationists should de-emphasize rigid enforcement of protected-area boundaries and absolute protection of conservation spaces from shifting people. Instead, conservationists should focus on developing durable means through which peopleconservationists, land managers, tourists, researchers, etc.can negotiate access to resources when this is both necessary and feasible without precluding potential future uses. The perceived imperatives of conservation reinforce the short-term focus of conservation practice. Land-use changes contributing to biodiversity loss have proceeded rapidly and widely in the last century. The rapidity of these changes in many areas has created the impression that unless significant efforts are made in the short term, insignificant levels of biodiversity will survive in the long term. Furthermore, regardless of the soundness of estimates or predictions of environmental change, short-term threats must logically be reduced if their persistence prevents the attainment of long-term goals. For instance, the short-term threat of human hunting must be reduced to ensure the long-term survival of chimpanzees, even though habitat loss is the primary long-term threat (Kormos & Boesch 2003). Altogether, current understanding of recent and

281 expected environmental changes creates substantial pressure to address and reduce perceived threats to biodiversity as quickly as possible. Lost in this situation is consideration of the longterm consequences of conservation interventions. Neglecting long-term planning in conservation practice because of the perceived urgency of short-term threats is tantamount to wasteful resource use, in which long-term sustainability is neglected because of perceived resource abundance. Partially as a result of the pressure to reduce short-term biodiversity threats, conservation practitioners frequently address perceived problems by using generalized strategies that have been successful elsewheresuch as IUCNs general strategy of establishing integral protection zones (see Chapter 2)without considering the appropriateness of these strategies in the specific contexts of particular protected areas. As a result, these strategies often represent blunt means of achieving conservation goals, perhaps achieving some degree of success in attaining specific goals, only while creating new problems because of their inappropriateness in a particular context. This dissertation argues that the application of IUCNs general strategy of establishing integral protection zones will likely fail long-term, if not short-term, conservation goals in Malis Bafing reserve, because the rigid and absolute boundaries this strategy creates are inappropriate for the biophysical and socioeconomic contexts of the Bafing area. There are many comparable cases in which generalized conservation strategies have proven clearly inappropriate for meeting conservation goals, and have been counterproductive to conservation success (e.g. Brockington & Homewood 1996; Daniels & Bassett 2002; Ite 2001; Koenig & Diarra 1998; Neumann 1997; Redford 1991). While there are certainly conservation strategies that are widely appropriate and not strongly influenced by local conditionsreducing hunting pressure, for instance, will reduce population decline for hunted animal species

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