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Mechanisms of Small Scale Foraging by Large African Herbivores

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Mechanisms of Small Scale Foraging by Large African Herbivores Satisfying Giant Appetites Mechanisms of small scale foraging by large African herbivores Yolanda Pretorius Promotores Prof. Dr. H.H.T. Prins Professor of Resource Ecology Prof. Dr. R. Slotow Professor of Biological Sciences, University of Kwa-Zulu Natal (South Africa) Co-promotor Dr. W.F. de Boer Lecturer at the Resource Ecology Group Thesis committee Dr.A.D. Rijnsdorp Wageningen University (Netherlands) Prof.Dr. S. Verhulst University of Groningen (Netherlands) Prof.Dr. H. Udo de Haes Leiden University (Netherlands) Dr. A.G. Toxopeus International Institute for Geo-information Science and Earth Observation Enschede (Netherlands) This research was conducted under the auspices of the C.T. de Wit Graduate School for Production Ecology and Resource Conservation Satisfying Giant Appetites Mechanisms of small scale foraging by large African herbivores Yolanda Pretorius Thesis submitted in partial fulfillment of the requirements for the degree of doctor at Wageningen University by the authority of the Rector Magnificus Prof. Dr. M.J. Kropff in the presence of the Thesis Committee appointed by the Doctorate Board to be defended in public on Monday 23 November 2009 at 11 AM in the Aula. Yolanda Pretorius Satisfying giant appetites: mechanisms of small scale foraging by large African herbivores, 141 pages. PhD-thesis, Wageningen University, Wageningen, the Netherlands (2009) with summaries in English, Dutch and Afrikaans. ISBN 978-90-8585-478-4 Preface The true meaning of the phrase ‘you are what you eat’ only became clear to me during the course of this study, except that for herbivores I think it should be ‘you are what you are constrained to eat’. This study started off rather broadly with the aim of determining the mechanisms behind small scale foraging by large herbivores. With guidance, trial and error I soon realized that a holistic approach leads to confusion and increases the chances of misinterpretation of results as it is impossible to account for all of the variability present, even at small scales. Eventually, after adopting a reductionist approach and using field experimentation as a tool to reduce variability, the study evolved to focus on how body mass differences determine mouth morphology, patch selection and daily diets of large African herbivores. I dedicate this thesis to my parents, Danie and Jorina and to the four mentors who have guided and inspired me throughout my academic career including my grandmother Drienie Smal, Jock McMillan, Paul Hendrik and Herbert Prins. This PhD journey has truly been an enriching experience both personally and in terms of my career. I hope you enjoy reading this work as much as I, despite occasional hardships, enjoyed creating it. Contents CHAPTER 1 General introduction………………………………………………………………. 1 CHAPTER 2 Why elephant have trunks, giraffe have long tongues & rhino have broad lips: mechanisms for how plants shape large herbivore mouth morphology ………….. 9 Y. Pretorius, K. Kortekaas, M. van Wijngaarden, W.F. de Boer, R. Slotow, & H.H.T. Prins CHAPTER 3 Soil nutrient status determines how elephant utilize trees and shape environments ……………………………………………………………………... 25 Y. Pretorius, W.F. de Boer, R. Slotow, C. van der Waal & H.H.T. Prins CHAPTER 4 Large herbivore responses to nutrient heterogeneity in an African savanna……… 39 Y. Pretorius, W.F. de Boer, M.B. Coughenour, H.J. de Knegt, H. de Kroon, C.C. Grant, I. Heitkonig, E. Kohi, E. Mwakiwa, M.J.S. Peel, A.K. Skidmore, R. Slotow, C. van der Waal, F. van Langevelde, S.E. van Wieren & H.H.T. Prins CHAPTER 5 Diet selection of African elephant over time shows changing optimization ……... 57 currency Y. Pretorius, J.D. Stigter, W. F. de Boer, S.E. van Wieren, C.B. de Jong, H.J. de Knegt, C.C. Grant, I. Heitkönig, N. Knox, E. Kohi, E. Mwakiwa, M.J.S. Peel, A.K. Skidmore, R. Slotow, C. van der Waal, F. van Langevelde, H.H.T. Prins CHAPTER 6 Synthesis: why size matters at small scales………………………………............. 77 Y. Pretorius References ……………………………………………………………………...... 101 Summary …………………………………………………………………............ 120 Samenvatting – Dutch Summary ………………………………………………... 124 Samevatting – Afrikaans Summary……………………………………………… 130 Affiliation of coauthors ………………………………………………………..... 135 Acknowledgements ……………………………………………………………... 136 Curriculum Vitae ……………………………………………………………....... 139 PE&RC PhD Education Certificate ……………………………………………... 141 Abstract Pretorius Y., 2009. Satisfying giant appetites: mechanisms of small scale foraging by large African herbivores Variation in body mass allows for resource partitioning and co-existence of different species. Body mass is also seen as the main factor governing nutrient requirements in herbivores as metabolic rate and requirements have often been found to scale to ¾ power of body mass. Although the consequences of body mass on foraging behaviour of herbivores has been extensively studied, the mechanism behind how body mass differences determines the small scale foraging patterns of especially larger herbivores, has up to now been unclear. In this study, I looked at how body mass and small scale vegetation characteristics shaped the mouth morphology of herbivores and how body mass of a herbivore affects the scale at which intake is maximized. More over, I looked at what spatial scales a mega-herbivore can select nutrient rich patches, and at the trade-offs between quality and quantity of plant species as forage to be included in the diet of different size herbivores. The results indicate that the dilution of plant mass and more specifically leaf mass in space requires that mega-herbivores have enlarged soft mouth parts to compensate for this dilution. In the analysis of patch selection by herbivores, I introduce the novel concept of nutrient load, which provides a way of expressing the total available nutrients to a herbivore per grain of a specific spatial scale. I show that solitary animals and/or animals with extreme mouth sizes are able to select the aggregation of patches with either the largest patch sizes or highest local nutrient concentrations which together yield the highest total nutrient loads at a spatial scale of 2500m2. Finally, I demonstrate, using linear programming techniques with multiple nutrients as constraints, how a mega-herbivore’s daily diet choice is determined by forage abundance whereas a small herbivore is more constrained by fibre. Keywords: patch selection, allometry, spatial scaling, body mass, mouth morphology, linear programming CHAPTER 1 GENERAL INTRODUCTION 1 Changes in abiotic environmental factors such as atmospheric temperature and moisture levels have been present since the origin of life. These changes occur at both a macro and micro scale leading to spatial and temporal heterogeneity in the occurrence of biotic elements. Organisms either go extinct or adapt to environmental changes through the process of natural selection, which is the basis of speciation and the principle by which variation in a trait that enhances survival and reproduction is preserved (Darwin 1859). Millions of years of evolution have selected the fitness maximization strategies that extant species possess. However, over a much shorter time, humans have heavily impacted on and artificially changed environmental conditions (Prins & Gordon 2008) so that extant species’ ability to adapt has been challenged to the extreme. For example, during the last two centuries Africa’s large herbivores were first threatened by extinction from uncontrolled hunting activities and poaching (Pringle 1982), followed by unprecedented habitat conversion for agricultural practices (Olff et al. 2002) only to be left with a future where the effects of global warming might present an even bigger challenge to survival. Grazing ecosystems are among the earth’s most endangered terrestrial habitats (Frank et al. 1998) and with increasing CO2 levels, C4 grasses, which dominate savannas, are expected to decrease (Bond et al. 2003) with concomitant effects on wild grazing species (Prins & Gordon 2008). Although predictions on the possible effects of global warming have received much attention during the last decade (Root et al. 2003 Google, cited 1038 times), the immediate threat of habitat loss through land-use change by humans is often neglected (Brooks et al. 2002 Google, cited 284 times). Nearly half the world’s vascular plant species and one third of terrestrial vertebrates are endemic to 25 biodiversity “hotspots”, which historically covered 12 % of the land’s surface but today less than 1.4 % of intact habitat remains (Brooks et al. 2002). The inevitable question is, with increasing human impact, will large herbivores be able to adapt and survive or will they face another mass extinction? To prevent such an extreme event we need to know how large herbivores satisfy their daily nutrient requirements and how they are adapted to do so. More over, prediction of the distribution of large herbivore species, either for the purpose of conservation or management, requires an understanding of the mechanisms behind resource use by these species. Historically, studies often focused on large scale habitat use and distribution of large African herbivores using a descriptive approach (McNaughton & Georgiadis 1986). However, small scale foraging processes can significantly affect these large scale distribution patterns (Shipley 2007). In this study, I have used an experimental approach to determine the mechanisms behind small scale foraging by large African herbivores. 2 Contribution to foraging ecology Optimal foraging theory,
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