An Ecological Reconnaissance of Wilpattu National Park, Ceylon
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JOHN F. EISENBE An Ecological and MELVYN LOCKHA Reconnaissance of Wilpattu National Park, Ceylon SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY NUMBER 101 SERIAL PUBLICATIONS OF THE SMITHSONIAN INSTITUTION The emphasis upon publications as a means of diffusing knowledge was expressed by the first Secretary of the Smithsonian Institution. In his formal plan for the Insti- tution, Joseph Henry articulated a program that included the following statement: "It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge." This keynote of basic research has been adhered to over the years in the issuance of thousands of titles in serial publications under the Smithsonian imprint, com- mencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the following active series: Smithsonian Annals of Flight Smithsonian Contributions to Anthropology Smithsonian Contributions to Astrophysics Smithsonian Contributions to Botany Smithsonian Contributions to the Earth Sciences Smithsonian Contributions to Paleobiology Smithsonian Contributions to Zoology Smithsonian Studies in History and Technology In these series, the Institution publishes original articles and monographs dealing with the research and collections of its several museums and offices and of profes- sional colleagues at other institutions of learning. These papers report newly acquired facts, synoptic interpretations of data, or original theory in specialized fields. These publications are distributed by mailing lists to libraries, laboratories, and other in- terested institutions and specialists throughout the world. Individual copies may be obtained from the Smithsonian Institution Press as long as stocks are available. S. DILLON RIPLEY Secretary Smithsonian Institution INTERNATIONAL BOOK YEAR • 1972 111 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY NUMBER 101 JohnF.Eisenberg An Ecological I -KM J J 11 . m Jvcconnaissance 01 Wilpattu National Park, Ceylon SMITHSONIAN INSTITUTION PRESS CITY OF WASHINGTON 1972 ABSTRACT Eisenberg, John F., and Melvyn Lockhart. An Ecological Reconnaissance of Wil- pattu National Park, Ceylon. Smithsonian Contributions to Zoology, number 101. 118 pages 76 figures, 16 tables. 1972.—The results of this study are based on a 14- month survey in Wilpattu National Park, Ceylon, covering an area of some 580 square kilometers in the lowland dry zone scrub jungle. Special attention was paid to the major ungulate species and data concerning abundance, seasons of reproduction, and feeding patterns are discussed. Although the elephant {Elephas maximus) exhibits the lowest numerical density, it accounts for the second highest biomass, exceeded in biomass only by the axis deer, which is also the most numerically abundant ungulate in the lowland dry zone of Ceylon. Special attention was paid to the role of the elephant in the dry zone ecosystem. Composition of social groupings, seasonal habitat utilization trends, general behavior patterns, and population structure are discussed for the elephant (Elephas maximus), water buffalo (Bubalus bubalis), axis deer (Axis axis), sambar (Cervus unicolor), muntjac (Muntiacus muntjac), wild swine (Sus scrofa), and mouse deer (Tragulus meminna). Certain aspects of the behavior of the major predators are also discussed, including the general biology of the golden jackal (Canis aureus) and the Ceylon leopard (Panthera pardus). Social organizations of the un- gulate species are compared with special attention given to the mode of antipredator behavior exhibited by ungulates. Habitat utilization trends and comments on com- petition and ecological separation are included together with an estimate of numeri- cal abundance and biomass for the major ungulate species. Recommendations for management of the park system are discussed. Official publication date is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. UNITED STATES GOVERNMENT PRINTING OFFICE WASHINGTON : 1972 For sale by the Superintendent of Document*, U.S. Government Printing Office Washington, D.C. 20402 • Price $1.60 (paper cover) Contents Page Introduction 1 Acknowledgments 2 The Environment 4 Physical Description of the Park 4 Climate 4 Flora 7 The Mammalian Fauna 10 The Elephant (Elephas maximus) 10 Description and Aging Criteria 10 Criteria for Age Classes 11 Criteria for Distinguishing the Sexes 12 Social Groupings 13 Distribution in the Park 17 Habitat Utilization 18 The Use of Tall Grass Villus 18 Short Grass Feeding 19 Trail Utilization During a Long March 20 Tree-Pushing Behavior 21 Description of River-Crossings 21 Elephant Sign in the Habitat 21 Concluding Remarks on Habitat Utilization 22 Summary of Elephant Behavior Patterns 23 Activity 23 Patterns of Social Interaction 23 Behavior During Agonistic Encounters 24 Antipredator Behavior 25 Female-Young Relationships 26 Sexual Behavior 26 Behavior Patterns of Different Age and Sex Classes 28 Interspecific Social Relations 28 Population Structure 29 The Water Buffalo (Bubalus bubalis) 30 Description and Aging Criteria 30 Social Groupings and Habitat Utilization 30 Distribution in the Park 30 Movements 32 Habitat Utilization 32 General Observations on Behavior 33 Population Structure and Dynamics 36 The Spotted Deer (Axis axis) 37 Social Groupings 37 Distribution in the Park 38 Habitat Utilization 39 Behavior Patterns 40 Activity Rhythms and Group Behavior 40 Communication Mechanisms 41 Patterns of Social Interaction 42 Male-Male Interactions 42 in Page Male-Female Behavior 42 Female-Female Behavior 43 Female with Young 43 Antipredator Behavior 44 Interspecific Relationships 45 Population Trends 45 Seasons of Birth and Mating 45 Mortality and Biomass 47 The Sambar (Cervus unicolor) 47 Description and Age Classes 47 Grouping Tendencies 48 Habitat Utilization 49 Social Behavior 49 Antipredator Behavior 50 Population Trends 50 The Muntjac or Barking Deer (Muntiacus muntjac) 51 The Mouse Deer (Tragulus meminna) 52 Summary Statement on the Small Forest Ungulates 54 The Wild Swine (Sus scrofa) 55 Description and Definition of Age and Sex Classes 55 Social Groupings 55 Reproduction 55 Distribution in the Park 57 Feeding 57 Habitat Utilization 57 Summary of Swine Behavior 57 Activity 58 Patterns of Social Interactions 58 Male-Male Interaction 59 Male-Female Behavior 59 Female-Female Behavior 59 Adult-Young Relationships 59 Interrelationships of Young Pigs 60 Antipredator Behavior 60 Interspecific Relationships 60 Population Trends 60 The Jackal (Cants aureus) 61 Description and Social Organization 61 Distribution in the Park 62 Behavior Patterns 62 Predation Behavior 63 Population Structure and Dynamics 63 The Sloth Bear (Melursus ursinus) 63 The Leopard (Panthera pardus) 65 Home Range 66 Activity 69 Stalking and Predation 69 Reproduction 70 Social Behavior 70 General Comments on Sociability 70 Communication Mechanisms 71 Scratching Trees 71 Urine Marking 71 Scrapes 71 Calling 71 Review of Social Roles 73 Biomass of Prey and Home Range Occupancy 74 Page The Smaller Carnivores 75 The Mongooses {Herpestes) 75 The Civets (Viverricula and Paradoxurus) 75 The Smaller Cats 75 The Primates and Other Mammals 75 The Primates 75 Comments on Other Mammals 77 Comments on the Nonmammalian Fauna of Wilpattu 78 The Avian Fauna 78 The Reptile Fauna 79 A Consideration of the Termites 79 Synthesis and Comparisons 81 Trends in the Evolution of Social Behavior as Exemplified by Ungulates 81 Review of Ungulate Evolution 81 Adaptation to Open Country 82 The Social Organizations of Ungulates 82 Ungulate Social Systems and Antipredator Behavior 83 The Muntjac or Barking Deer 83 The Wild Swine 84 The Axis Deer 84 The Water Buffalo 85 The Asiatic Elephant 86 Relationship of Social Organization to Antipredator Behavior 87 Communication Mechanisms and the Evolution of Social Organizations ... 88 Summary of Other Social Mechanisms Manifest in Ungulate Groupings 89 Trends in Habitat Utilization 90 Competition and Ecological Separation 98 Numerical Abundance and Biomass 101 Recommendations for Management 104 Literature Cited 107 Appendix A: Methods Employed in Censusing 110 Appendix B: List of Some Dominant Plants of the Lowland Dry Zone 115 Index 117 John F. Eisenberg An Ecological and Melvyn Lockhart Reconnaissance of Wilpattu National Park, Ceylon Introduction Although the study here reported was addressed pri- actual discussion of the methods employed, see Appen- marily to the problem of conserving Ceylon's elephant dix A. population, it has been necessary to survey all the ma- A map of Wilpattu National Park and the surround- jor ungulate populations in the National Park in order ing sanctuary and intermediate zones is portrayed in to assess such subtle effects as competition and cooper- Figure 1. In our survey, we intensively studied the ation among the various populations of large mam- West Sanctuary, parts of the North Sanctuary, and mals. Special attention was directed towards censusing the major park area including the old south interme- the populations of large ungulates resident in the parks diate zone which last year was included in the park and to gather data concerning the seasonal movement itself. Two camps were established: a temporary camp patterns with respect to the distribution of food and on the banks of the Moderagam Am, which forms the water. Methodology was carried out in a classic pat- north boundary of the park, and a permanent camp tern. Two types