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Library of Congress Cataloging-in-Publication Data
Elephants and savanna woodland ecosystems : a study from Chobe National Park, Botswana / edited by Christina Skarpe, Johan T. du Toit and Stein R. Moe.
pages cm
Includes bibliographical references and index.
ISBN 978-0-470-67176-4 (cloth)
1. Elephants– Botswana– Chobe National Park. 2. Grassland ecology– Botswana– Chobe National Park. 3. Chobe National Park (Botswana) I. Skarpe, Christina, 1946- editor of compilation. II. Du Toit, Johan T., editor of compilation. III. Moe, Stein R., 1960- editor of compilation.
QL737.P98E443 2014
599.67096883– dc23
2013046029
A catalogue record for this book is available from the British Library.
Wiley also publishes its books in a variety of electronic formats. Some content that appears in print may not be available in electronic books.
Cover image: Front cover: Elephants Loxodonta africana browsing on Croton megalobotrys.
Photo: Stein R. Moe.
Back cover: Alerted impala ram and a group of bull elephants, Mababe, Botswana.
Photo: Christina Skarpe.
Cover design by Design Deluxe
1 2014
After a long journey through the dry Kalahari sand woodlands stretching over northern Botswana, a spectacular sight confronts one on arrival at the Chobe River. The emerald landscape of meadows and water on the associated floodplain is thronged by elephants, buffaloes and hippos, alongside numerous smaller ungulates and birds. This is a prime example of a megaherbivore-dominated ecosystem, lacking only the rhinos that were once also there. However, the very largest of these herbivores has been disrupting the structure and diversity of the riparian woodland, to the consternation of wildlife managers, tourist operators and visitors. The Chobe River front has attained notoriety for the woodland devastation wrought by the elephant concentrations there. Gaunt trunks of dead trees stand amongst battered shrubs above a sparse herbaceous cover. Should not something be done about the elephants to rectify this situation? Along with Tsavo National Park in Kenya, the state of woodland destruction at Chobe is commonly invoked as justification for culling elephants to alleviate the vegetation transformation and its ramifications for biodiversity.
This was the context for the BONIC project, established as an institutional collaboration involving the Norwegian Institute for Nature Research, the Norwegian University of Life Sciences, and the Botswana Department of Wildlife and National Parks. Its aim was to advance local capacity to address this and other problems in the management of Botswana's rich wildlife resource, through research and training. The expectation was that participating wildlife ecologists from the far north, untainted by the polarised standpoints about the management of elephants that prevail within African countries, would be openly receptive to the prospect of culling elephants if ecologically justified. After all, Norway has persisted in harvesting of whales, despite international criticism. The crucial research needed was to interpret the impacts that elephants were obviously having on riparian woodlands within the context of other ecosystem components and processes. Hence studies were focused not merely on the elephants, but also on their effects on soil properties and tree regeneration, the consequences of woodland transformation for other browsers and grazers, and ramifying influences on predators, small mammals and even some groups of birds. The chapters of this book document the findings from this comprehensive suite of studies, summarising or elaborating numerous papers that have appeared in scientific journals as well as the contents of several unpublished theses.
Surprisingly to some, these studies did not find adverse consequences of the obvious elephant impacts on vegetation for any animal species, apart from bushbuck, which have declined in abundance from earlier times when the bush cover was much thicker. The stumbling block for restoration of the woodlands lies in the high local abundance of impala, a much smaller herbivore that is a mixed grazer-browser. Exclosure plots demonstrated that their browsing of tree seedlings is so thorough that very few of these seedlings have much chance of escaping towards tree height. Moreover, rather than competing for browse, elephants favour different woody species from those utilized by browsing ruminants. This finding reinforces the suggestion that megaherbivore extinctions largely through human hunting had negative consequences for post-Pleistocene large mammal diversity worldwide (Owen-Smith, 1987, 1989). The disturbing impacts that mammoths, mastodons and other very large herbivores must have had on woodlands and forests would have brought more browse within the reach of smaller browsers, while increasing the extent of meadows for grazers. Hence the largest herbivores can facilitate the coexistence of smaller species through these mechanisms, rather than competing with them for shared vegetation resources.
Nevertheless, the extent of the woodland destruction along the Chobe River front is regrettable negative scenically. However, happenings in the more distant past have probably contributed to this situation. After the extirpation of the elephants, browsing antelope became decimated by rinderpest, allowing unfettered growth by trees. Are the Chobe woodlands merely reverting to the messy state that had prevailed when herbivores from elephants size down had all been hugely abundant? But expanded human pressures are an exacerbating influence, funnelling elephants into a narrow section of river front between Kasane town and villages within the Kachikau enclave, and blocking movements across the international border into Namibia.
Fundamental questions remain about how riparian woodlands can withstand the elephant concentrations that develop along Africa's “pristine” rivers during the dry season. Seasonal elephant densities even greater than those near the Chobe River have become established along the Linyanti River to the west, following the drying of the Savuti Channel and other water sources in the interior. Will the vegetation transformation there progress towards the state prevalent along the Chobe? Or be averted by the wider scope that elephants have for movement in the Linyanti region? Research is in progress to assess this trend and where it might eventually lead.
Elephants and their impacts on trees have drawn most of my attention. But the chapters in this book encompassing broader ecosystem ecology will be more widely valuable as a counterpoint to the renowned studies undertaken in the grassland ecosystem prevalent in the Serengeti region of Tanzania. Most of Africa is very different in aspect from Serengeti, having vast areas of fairly well wooded savanna occupying nutrient deficient soils of granitic or Aeolian origin. In these parts the very largest grazers and browsers assume dominance of the herbivore biomass rather than the “plains game” typical of the East African plateau. Botswana is an amazingly diverse country, with dry savanna woodlands juxtaposed with the wetlands of the Okavango Delta and abutting Kalahari semi-desert in the south. Interspersed are localities where the concentration of herbivores and predators rival those in the Serengeti ecosystem. There are huge challenges in managing Botswana's rich wildlife legacy, not least because of the continuing expansion of the elephant population, approaching 150,000 animals at the time of writing. A major contribution of the BONIC programme was its fostering of local wildlife scientists equipped with the qualifications to take responsibility for this custodianship. Regional planning is well advanced for Chobe National Park to become a component of the vast Kavango-Zambezi Transfrontier Conservation Area stretching from Botswana through adjoining parts of Namibia, Zimbabwe, Zambia and Angola. The findings from the BONIC studies will help inform this ambitious development in its aim of promoting the coexistence of people and wildlife.
The common image of an African savanna, held by people living far from savanna environments, is a landscape of short-cropped grass and scattered Acacia trees that is teeming with medium-sized grazing ungulates such as plains zebra, Equus quagga, and blue wildebeest, Connochaetes taurinus. This is an image based largely on the Serengeti-Mara ecosystem in Tanzania and Kenya, well known from extensive popular science publications, films and TV programs, as well as seminal scientific publications. The woodland savannas of Chobe National Park in Botswana, the focus for this book, present a very different image. Whereas mean annual rainfall is about the same in large areas of Serengeti and Chobe, most other ecological conditions are different. Instead of the nutrient-rich volcanic soils of the Serengeti savannas, the Chobe woodlands grow on nutrient-deficient Kalahari sand and instead of scattered fine-leafed Acacia trees, the Chobe woodlands consist primarily of large, broad-leafed trees. The abundant medium-sized grazers in the Serengeti are replaced in the Chobe woodlands by a dominance of larger-bodied species such as African buffalo, Syncerus caffer, and elephant, Loxodonta africana, with more than half the herbivore biomass in Chobe National Park contributed by elephants alone.
Elephants were virtually exterminated from the Chobe ecosystem by an intense bout of commercial ivory hunting in the late 19th century. Over the following decades, woodlands established on the previously open narrow strip with alluvial soil close to the Chobe River. These woodlands were different from those on the Kalahari sand in most of the Chobe National Park. Once the elephant population eventually began recovering, elephants killed the trees in these newly established woodlands by debarking them, and since the 1960s managers and conservationists have been concerned about the destruction of the scenic woodlands along the Chobe River. This ‘Chobe elephant problem’ was the rationale for the Botswana–Norway Institutional Cooperation Project (BONIC), a research and capacity building project run in cooperation between the Botswana Department of Wildlife and National Parks (DWNP) and two Norwegian research institutions: the Norwegian Institute for Nature Research (NINA) and the Norwegian University for Life Sciences (UMB), with funding from the governments of Botswana and Norway. BONIC operated from 1997 to 2003 and encompassed studies on diverse aspects of the Chobe ecosystem, all with a focus on the ecological implications of the increasing elephant population. At the termination of the project, a workshop was held in Kasane, adjacent to the study area, over 13–15 March 2003 including about 50 people from DWNP, NINA, UMB, University of Botswana, some non-governmental organisations and three specially invited experts: Patrick Duncan, Norman Owen-Smith and Anthony R.E. Sinclair. The workshop resulted in the first compilation of preliminary results from the project in a volume of proceedings, edited by Mark Vandewalle and published by the Government Printer, Gaborone, Botswana in 2003. It was followed in 2004 by an overview article about the project in the journal AMBIO by C. Skarpe and 26 co-authors from the project.
The workshop in 2003 gave rise to the idea of an edited book drawing from the main results from the project. First, over the ten years following the completion of the project, the results of various sub-projects were written up as graduate theses and peer-reviewed journal publications. Finally, this book synthesizes the ecological research conducted under the auspices of BONIC. The book's aim is to present results from the project related to the effects of elephants on ecosystem dynamics and heterogeneity, and finally to discuss the extent to which there is an ‘elephant problem’ in Chobe. The book compiles information from a nutrient-poor and elephant-rich savanna to allow comparison with other African savannas, for example: the nutrient-rich Serengeti-Mara ecosystem as presented in the three books edited by A.R.E. Sinclair and colleagues (University of Chicago Press 1979, 1995, 2008); the Kruger National Park on mixed soil types, as described in the book edited by J.T. du Toit, K.H. Rogers and H.C. Biggs (Island Press 2003); the nutrient-poor and elephant-free Nylsvley savanna described by R.J. Scholes and B.H. Walker (Cambridge University Press, 1993).
Most studies in the BONIC project are included in this book, some constituting individual chapters. We refer as far as possible to published data from the project, although data from PhD and MSc theses are also referred to along with some previously unpublished data, which are included without references. Four staff members from DWNP completed their PhD projects within BONIC and all of them contributed as authors to this book: Shimane W. Makhabu, Gaseitsiwe S. Masunga, Lucas P. Rutina and Cyril L. Taolo. Eight staff members from DWNP completed their MSc studies within BONIC: Kingsley M. Leu, Itani Mathumo, Thato B. Morule, David K. Mosugelo, Sekgowa S. Motsumi, Elsie T. Mvimi, Gosiame Neo-Mahupeleng and Claudia S. Zune. Their work has directly and indirectly contributed to this book.
Neither the BONIC project nor this book could have come into existence without DWNP, being the project leader, and the Ministry of Environment, Wildlife and Tourism of Botswana and their staff. Apart from staff members appearing as authors in the book, we particularly thank Jan Broekhuis, Joe Matlhare, Sedia Modise, Dan Mughogho, Bolt Othomile and Botshabelo Othusitse. Further, Thatayaone Dimakatso, Frederick Dipotso, Wilson Marokane, Moses Mari, Mpho Ramotadima, Ditshoswane Modise (now deceased), Zenzele Mpofu (now deceased), Lettie Sechele and many others helped as counterparts and field assistants in data collection and research. Abraham Modo (now deceased), then District Coordinator for Ngamiland, contributed by allowing his staff to participate in the project as field assistants.
The Norwegian Embassy in Gaborone provided valuable support to the project, and we are particularly indebted to Jan Arne Munkeby, who was the Norwegian Chargé d'Affaires at the time of project inception. Britt Hilde Kjølås, Embassy Secretary, also provided invaluable assistance during the initial phase of the project.
The chapters of this book were improved by the critical comments of independent expert reviewers and the following are thanked for their valuable contributions: George Batzli, Jane Carruthers, David H. M. Cumming, Hervé Fritz, Jacob R. Goheen, Ricardo Holdo, R. Norman Owen-Smith, Steward T.A. Pickett, Robert J. Scholes, Peter Scogings, Anthony R.E. Sinclair, Izak Smit, Marion Valeix and George Wittemyer. We sincerely thank the editing team at Wiley-Blackwell, particularly Ward Cooper, Kelvin Matthews and Carys Williams for encouraging and efficient collaboration as well as great patience. We further thank Ola Diserud and Andreas Brodén for contributions to Chapters 5 and 9, respectively; Marit Hjeljord for drawing many of the figures; and Lin Cassidy for drawing the map in Figure 1.3.
Finally, the entire text of this book was copyedited by Peter Frost who applied his writing skills, understanding of the English language, broad knowledge of African savannas and, above all, his meticulous professionalism, to substantially enhance the final product. For his valued contributions the editors owe Peter a large debt in gratitude.
The basis for this book was laid by ivory hunters and cattle herders operating more than a 100 years ago south of the Chobe River in what today is northern Botswana. In the book we explore how the virtual extinction of the elephants, Loxodonta africana, by ivory hunters in the second half of the 19th century and the simultaneous reduction in many ungulate populations caused by rinderpest, a viral disease spread with cattle, initiated a dramatic history of ecosystem perturbations, many of which are still ongoing. The dynamics involved many aspects of the ecosystem, quantitative as well as qualitative, including plants, soil and animals varying in multiple spatial and temporal scales. To analyse this complex heterogeneity, we adopted the framework presented by Pickett et al. (2003). A framework or a model is a simplification and generalisation providing a context, in this case allowing us to identify origin, spatial and temporal scale and pattern of variation. The framework is hierarchical in nature, allowing shifting between scales to assess different degrees of detail.
For a pattern of heterogeneity to exist in time and space there must be a substrate exhibiting the heterogeneity and an agent creating the heterogeneity by its actions on the substrate. For the pattern to be functionally meaningful there must also be some organism or process that responds to the pattern of the substrate, and generally there are also one or more factors controlling the interactions between agent, substrate and responder (Pickett et al., 2003). Therefore, to understand the function of ecological heterogeneity in a certain ecosystem, we need to specify what substrate exhibits the variation, what agent works on the substrate to cause the variation, what organisms or processes respond to the variation in the substrate and what factors control the interactions (Pickett et al., 2003; Figure 1.1). For example, on the alluvial soil along the Chobe River the increasing population of elephants, functioning as agents, have altered the vegetation, the substrate, by debarking and killing most of the large trees. The resulting tree-less state of the vegetation is being maintained in turn by seedling-eating impala, Aepyceros melampus, which are acting as controllers by preventing regeneration of the trees (Moe et al., 2009; Chapter 10). The change in the state of the substrate has led to adjustments in population size and/or behaviour of a number of herbivore and carnivore species, which thus act as responders. So have the increasing openness of the vegetation resulting from the elephant and impala activities been a disadvantage for the thicket-preferring bushbuck, Tragelaphus scriptus, but might have favoured the puku, Kobus vardonii, preferring open plains (Dipotso and Skarpe, 2006; Dipotso et al., 2007; Chapter 13).
Figure 1.1 The conceptual model of the Chobe ecosystem dynamics. Agents, for example, elephants, create or maintain heterogeneity on specific scales by acting on a substrate, in our example, the vegetation. Substrates change or are maintained in a certain state by action of the agent (or the absence thereof); in our example the vegetation changed from woodland (state 1) to shrubland (state 2). Controllers affect the action of the agent on a substrate or the resultant transition (and reversibility) between states of a substrate, for example, impala preventing regeneration of the woodland once elephants have killed the large trees. Responders are the variables influenced by the change or state transition of the substrate, for example, other herbivores. Drawing by Marit Hjeljord.
Agents such as elephants can create, maintain or transform states of the substrate upon which they are acting. They are agents of change only as long as they themselves change, for example in density or behaviour. When the substrate has ceased exhibiting directed change, the role of the agent may be to maintain the acquired state (Skarpe, 1992; Pickett et al., 2003). In the Chobe situation, the earliest mentioning of the alluvial flats above the flood plain, where the elephants now have killed the large trees, described a very open vegetation (Selous, 1881), which have been maintained in that state by the activities of large herbivores acting as agents and controllers (Chapters 4 and 12). A change in the agent and controller, such as the decline of elephants and ungulates following the ivory hunt and the rinderpest panzootic, respectively, initiated profound changes in the substrate by providing a window of opportunity for the establishment on the alluvial flats of the woodlands, which were later reduced with the recovery of the elephants constituting an agent of change (Chapters 4 and 10).
The heterogeneity framework described can be seen as a hierarchy of spatial and temporal scales, where small-scale pattern and processes may be contained within a such larger scale. Biotic and abiotic agents create heterogeneity in all ecosystems and at all temporal and spatial scales, from global climate governing continental-scale variation in plant and animal communities over decades, centuries and millennia to selective grazing by buffalo, Syncerus caffer, creating patchiness in grassland vegetation, termites modifying soil around their nest or twig-browsing greater kudu, Tragelaphus strepsiceros, causing differentiation of shoot growth within a tree canopy. Such differences in scale mean that an agent in one scale can be a responder or a controller at another, or sometimes have different roles at the same scale (Pickett et al., 2003). In a local scale on the Chobe flood plains buffalo are agents, increasing heterogeneity in the grass sward by creating and maintaining grazing lawns. Responders to this change in the substrate are other grazing species such as impala and puku utilising the lawns. In another scale buffalos are responders facilitated by elephant-induced changes in the vegetation, and controllers influencing the spatial distribution of elephant grazing (Taolo, 2003; Chapter 11). Other examples of multiple roles are the few remaining live trees on the alluvial flats. These trees are the substrate for elephant activity, but are also agents themselves in a smaller scale, causing local changes in soil properties and microclimate to which many organisms, plants and animals, respond (Campbell et al., 1994; Pickett et al., 2003). The book is mainly structured according to a spatial and temporal scale to which many human activities such as conservation and management relate, where vegetation is the main responder and elephants the agent creating, modifying or maintaining heterogeneity.