Ecological Studies. Analysis and Synthesis

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1 Ecological Studies Analysis and Synthesis Edited by W. D. Billings, Durham (USA) F. Golley, Athens (USA) O. L. Lange, Wiirzburg (FRG) J. S. Olson, Oak Ridge (USA) H. Remmert, Marburg (FRG) Volume 42

2 Ecology of Tropical Savannas Edited by B. J. Huntley and B. H. Walker With 262 Figures Springer-Verlag Berlin Heidelberg New York 1982

3 BRIAN J. HUNTLEY CSIR, Cooperative Scientific Programmes P.O. Box 395, Pretoria 0001, BRIAN H. WALKER Department of Botany and Microbiology University of the Witwatersrand P.O. Box 1176, Johannesburg 2000, ISBN -13: e-isbn -13: DOl: / Library of Congress Cataloging in Publication Data. Main entry under title: Ecology of tropical savannas. (Ecological studies; v. 42) Bibliography: p. Includes index. 1. Savanna ecology - Tropics. I. Huntley, B.l. (Brian 1.), II. Walker, B.H. (Brian Harrison), III. Series. QH84.5.E '2643' This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically those of translation, reprinting, re-use of illustrations, broadcasting, reproduction by photocopying machine or similar means, and storage in data banks. Under 54 of the German Copyright Law where copies are made for other than private use, a fee is payable to "Verwertungsgesellschaft Wort" Munich. by Springer-Verlag Berlin Heidelberg 1982 Sof tcover reprint of the hardcover 1 at edition 1982 The use of registered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. 2131/

4 Acknowledgements The symposium and workshop from which this volume proceeds was convened by the n Savanna Ecosystem Project, one of several large interdisciplinary cooperative research projects of the National Programme for Environmental Sciences. The support of the Council for Scientific and Industrial Research, the Department of Environmental Affairs, the Transvaal Provincial Administration, the Department of Agriculture and Fisheries and the National Parks Board is gratefully acknowledged. The preparation of camera-ready material for the publication of this book would not have been possible without the meticulous typing and endless perseverance of Marie Breitenbach and Margaret Orton. Our editorial task has been considerably lightened by Suzanne Winkelman and Shirley Ellis. Thanks are also due to the contributing authors, whose active participation in the meeting discussions and subsequent revision of their papers has helped provide an integrated account of the savannas of the southern continents. Editorial Note The inclusion of accent marks in Portuguese, French and Spanish words has not been possible due to the limitations set by the word processing system used in the preparation of the camera-ready copy.

5 Contents Section 1. Introduction Section 2. Structure. 3 I. Australian Savannas. J. WALKER and A. N. GILLISON 2. Brazilian "Savannas". G. EITEN Chaco and Caatinga - South American Arid Savannas, Woodlands and Thickets. E. H. BUCHER The Structure and Dynamics of a West African Savanna J. C. MENAUT and J. CESAR Southern African Savannas. B. J. HUNTLEY Woody Plant Biomass Distribution in Burkea africana Savannas M. C. RUTHERFORD Section 3. Determinants The Influence of Soils, Geomorphology and Geology on the Distribution of Plant Communities in Savanna Ecosystems. M. M. COLE The Influence of Soil Moisture Balance on Ecosystem Patterns in Southern Africa. K. L. TINLEY The Effect of Soil Nutrient Availability on Community Structure in African Ecosystems. R. H. V. BELL The Influence of Large Herbivores on Savanna Structure in Africa D. H. M. CUMMING Fire in Australian Tropical Savannas. C. J. LACEY, J. WALKER, and I. R. NOBLE Ecological Effect of Fire in Brazilian Cerrado. L. M. COUTINHO Ecological Effects of Fire in n Savannas. W. S. W. TROLLOPE 292 Section 4. Function " Physiological Ecology of Neotropical Savanna Plants. E. MEDINA Water Relations of Southern African Savannas G. C. BATE, P. R. FURNISS, and P. G. PENDLE

6 VIII 16. Factors Influencing the Consumption of Plant Products by Large Herbivores. N. OWEN-SMITH Response of the Humid Subtropical Grassland of to Defoliation. N. M. TAINTON Consumption and Decomposition in Tropical Grassland Ecosystems at Lamto, Ivory Coast. M. LAMOTTE Section 5. Nylsvley, a n Savanna Structure of the Nylsvley Savanna. B. J. HUNTLEY and J. W. MORRIS Towards the Development of an Energy Budget for a Savanna Ecosystem J. M. DE JAGER and T. D. HARRISON Phytomass, Seasonal Phenology and Photosynthetic Studies C. F. CRESSWELL, PAM FERRAR, J. O. GRUNOW, D. GROSSMANN, M. C. RUTHERFORD, and J. J. P. VAN WYK Nitrogen in the Burkea Savanna. G. C. BATE and CHARLOTTE GUNTON Trophic Ecology and Plant/Herbivore Energetics. M. V. GANDAR Litter Decomposition. J. W. MORRIS, J. J. BEZUIDENHOUT, and P. R. FURNISS Section 6. Dynamics and Management Aspects of the Stability and Resilience of Savanna Ecosystems B. H. WALKER and I. NoY-MEIR Stability of Plant-Herbivore Models and Possible Applications to Savanna. I. NoY-MEIR A Developing Computer Model of the Auob River Ecosystem. Kalahari Gemsbok National Park. A. M. STARFIELD, S. M. SHAPIRO, P. R. FURNISS, M. SEARS, P. F. RETIEF, P. T. VAN DER WALT, and M. G. L. MILLS Management Strategies for the Utilization of Southern African Savanna D. L. BARNES Section 7. Conclusion: Characteristic Features of Tropical Savannas 657 Subject Index

7 Contributors BARNES, D. L. BATE, G. C. BELL, R. H. V. BEZUIDENHOUT, J. J. BUCHER, E. H. CESAR, J. COLE, MONICA M. COUTINHO, L. M. CRESSWELL, C. F. CUMMING, D. H. M. DE JAGER, J. M. EITEN, G. FERRAR, PAM FURNISS, P. R. 21 Theron Street, De Bruin Park, Ermelo 2350, South Africa Department of Botany, University of Port Elizabeth, P.O. Box 1600, Port Elizabeth 6000, Kasungu National Park, P.O. Box 43, Kasungu, Malawi Department of Microbiology, Faculty of Agriculture, University of Pretoria, Pretoria 0002, Centro de Zoologia Aplicada, Universidad de Cordoba, Casilla de Correas 122, 500 Cordoba, Argentina Ecole Normale Superieure, Laboratoire de Zoologie, 46 Rue d'ulm, Paris 05, France Department of Geography, Bedford College, University of London, Regent's Park, London NWI 4NS, Great Britain Departamento de Ecologia, Instituto de Biociencias, Universidade de Sao Paulo, Sao Paulo, Brazil 2000, Department National Parks and Wildlife Management, P.O. Box 8365, Causeway, Salisbury, Zimbabwe Department of Agrometeorology, University of the Orange Free State, P.O. Box 339, Bloemfontein 9300, Veg-I B, Universidade de Brasilia, Brasilia DF, Brazil 2000, Langwell House, Willow Grove, Chislehurst, Kent BR 7 5BN, Great Britain

8 x GANDAR, M. V. GILLISON, A. N. GRUNOW, J.O. GUNTON, CHARLOTTE HARRISON, T. D. HUNTLEY, B. J. LACEY, c.j. LAMOTTE, M. MEDINA, E. MENAUT, J. C. MILLS, M. G. L. MORRIS, J. W. NOBLE, 1. R. NoY-MEIR,1. OWEN-SMITH, N. PENDLE, P.G. Institute for Natural Resources, University of Natal, P.O. Box 375, Pietermaritzburg 3200, Division of Land Use Research, CSIRO, P.O. Box 1600, Canberra City, ACT 2601, Australia Department of Plant Production, University of Pretoria, Pretoria 0002, 2000, Department of Agrometeorology, University of the Orange Free State, P.O. Box 339, Bloemfontein 9300, Cooperative Scientific Programmes, CSIR, P.O. Box 395, Pretoria 0001, Division of Land Use Research, CSIRO, P.O. Box 1666, Canberra City ACT 2601, Australia L'Ecole Normale Superieure, Laboratoire de Zoologie, 46 Rue d'ulm, Paris 05 France IVIC, Centro de Ecologia, Apartado No 1827, Caracas 10 1, Venezuela Ecole Normale Superieure, Laboratoire de Zoologie, 46 Rue d'ulm, Paris 05, France Kalahari Gemsbok National Park, Private Bag X5890, Upington 8800, Datametrical Services, Department of Agriculture and Fisheries, Private Bag X116, Pretoria 0001, Department of Environmental Biology, Research School of Biological Sciences, Australian National University, P.O. Box 475, Canberra City ACT 2601, Australia Department of Botany, Hebrew University of Jerusalem, Jerusalem, Israel Centre for Resource Ecology, University of the Witwatersrand, P.O. Box 1176, Johannesburg 2000, South Africa 2000,

9 XI RETIEF, P. F. RUTHERFORD, M. C. SEARS, M. SHAPIRO, S. M. STARFIELD, A. M. TAINTON, N.M. TINLEY, K. L. TROLLOPE, W. S. W. VAN DER WALT, P. T. VAN WYK, J. J. P. WALKER, B. H. WALKER, J. National Parks Board, Private Bag X404, Skukuza 1350, Botanical Research Institute, Private Bag X101, Pretoria 0001, Department of Applied Mathematics, University of the Witwatersrand, P.O. Box 1176, Johannesburg 2000, Department of Applied Mathematics, University of the Witwatersrand, P.O. Box 1176, Johannesburg 2000, Department of Civil and Mineral Engineering, University of Minnesota, 112 Mine and Metallurgy, 221 Church Street SE, Minneapolis, MN 55455, USA Department of Pasture Science, University of Natal, P.O. Box 375, Pietermaritzburg 3200, Blue Bend, P.O. Box 2021, Beacon Bay 5205, South Africa Department Agronomy, Faculty of Agriculture, University of Fort Hare, Private Bag X13l4, Alice 5700, National Parks Board, Private Bag X404, Skukuza 1350, Department of Botany, Potchefstroom University for CRE, Potchefstroom 2520, 2000, Division of Land Use Research, CSIRO, P.O. Box 1666, Canberra City ACT 2601, Australia

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