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Amphibian Diversity and Breeding Behaviour in the Okavango Delta

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Amphibian Diversity and Breeding Behaviour in the Okavango Delta Amphibian diversity and breeding behaviour in the Okavango Delta M. le Roux 21251304 Dissertation submitted in fulfilment of the requirements for the degree Master of Science at the Potchefstroom Campus of the North-West University Supervisor: Prof. L.H. du Preez Co-supervisor: Dr. C. Weldon November, 2010 PREFACE AND ACKNOWLEDGEMENTS This project was made possible by the fact that I was under full time employment in the Okavango Delta, working for the safari operator Okavango Wilderness Safaris. This offered me the opportunity not only to be based out of my study region and conduct my research on a part time basis, but also allowed much time to understand the ecosystem and constantly observe changes in my surrounding environment. During the course of the project, I was first based in Xigera (from Feb 2007 to Feb 2009) and then at Mombo (from Mar 2009 to Jul 2010). Therefore, in addition to the official field trips that were arranged, additional, ad hoc observations were possible outside of these official trips; an opportunity which proved invaluable, especially to Chapter 3. The nature of employment was such that I worked seven days a week for three months at a time, never leaving the safari lodge area, followed by a one month break outside of the Okavango Delta. Although being stationed in the study region for the duration of the study allowed for a thorough investigation of all the aims, full time employment did have certain drawbacks that will be pointed out where necessary in each chapter, the most prominent of which was the difficulty in standardisation of sampling sites and times. I would like to thank my supervisors, Prof. Louis du Preez and Dr. Ché Weldon, for their constant support, guidance and enthusiasm throughout the course of this project, and for their review and constructive criticism during the creation of this document. Their knowledge and expertise in the field of amphibians is remarkable and my goals would not have been accomplished without their assistance. I am indebted to the numerous field assistants who have accompanied me and humoured me during the many field trips and countless hours spent searching for frogs in the Okavango Delta waters: Simon Byron, Kai Collins, Tony Reumerman, Anton & Carrie Wessels, Kgabiso Lehare, Brian Rhode, Chantelle Venter, Glynis Humphrey, Gerhard du Preez, Pete Myburg and Katie Walker. I wish to thank Okavango Wilderness Safaris and their Environmental Department in particular their assistance during the project, for access to their concessions and lodges where the research was conducted, use of their vehicles and equipment, and flexibility with regards to time granted for my studies. To everyone involved, especially: Brandon & Deborah Kemp-White, Map Ives, Kai Collins, Anton Wessels and Tapera Sithole. Also, I would like to thank the Wilderness Wildlife Trust for the funding they provided to make this project possible, specifically: Russell Friedman and Chris Roche. ii To the many others who have assisted me in various ways, I express gratitude: AACRG (African Amphibian Conservation and Research Group) for materials, literature and support; HOORC (Harry Oppenheimer Okavango Research Centre) for facilities and assistance, particularly Lars Ramberg, Cornelis vanderPost and Mike Murray-Hudson; Pretoria National Zoological Gardens; John Mendelsohn; Dirk Cilliers; Stefan & Francis Siebert; Adrian Haagner; Jaco Bezuidenhout; Suria Ellis; Carina de Beer; Sven Bourquin and David Parry. For their ongoing trust, support and encouragement: Simon Byron, Bryan & Marna Williams, Gerhard, Lene & Anneke le Roux, Martin & Kylie Farrell; Charles, Maryke, Jerome and Catherine Byron. As well as anyone else that may have been accidentally left out. Hyperolius parallelus from the Okavango Delta iii ABBREVIATIONS USED IN THE TEXT Bd Batrachochytrium dendrobatidis cm centimetre(s) g gram(s) l litre(s) m metre(s) mg milligram(s) ml millilitre(s) mm millimetre(s) QDGC quarter degree grid cell EXPLANATION OF NON-ENGLISH TERMS Vlei Part of a watercourse which spreads out over a flat, depressed valley characterised by specialised water-based vegetation iv ABSTRACT Amphibians are of great ecological importance and a loss of species will have widespread and dire consequences. Recent population declines and extinctions have resulted in amphibians being labelled the most threatened vertebrate class on a global scale. The unique Okavango ecosystem is well known and documented, yet the amphibians of this region are poorly known. This project aimed at assessing diversity in the Okavango Delta by testing isolation as a possible driver for community composition; determining the effect of hydrology on breeding behaviour; and assessing the status and prevalence of the pathogen Batrachochytrium dendrobatidis (Bd) responsible for the widespread epidemic chytridiomycosis implicated in amphibian decline. Using various monitoring techniques, observations of species occurrence were made at three locations representing different degrees of isolation over a 20 month period. Breeding indicators were observed and frogs were screened for amphibian chytrid fungus. A total of 29 species were recorded, and results indicated that there were no significant differences in community composition between the sampled localities. Species presence, however, was significantly correlated with habitat type. Thus, the availability of suitable habitat appears to be driving amphibian diversity patterns, rather than geographic isolation; and increased habitat diversity near the Delta periphery explains increased amphibian diversity in these areas. Results from breeding indicators suggested that reproduction in continuous breeders was correlated with the annual flood as well as rainfall, whilst that of explosive breeders was correlated with rainfall alone. It is thus proposed that opportunistic breeding behaviour for some amphibian species is driven by the hydrology of the ecosystem; and this may be explained by increased biological production associated with the flood pulse. Outcomes highlight the unique nature of the Okavango Delta system, and emphasises the need for its preservation. A total of 249 swab samples were collected and screened for amphibian chytrid fungus. The geographical distribution of collection samples were evenly spread throughout the localities, and were obtained from at least 25 amphibian species. Analyses proved negative for Bd for the 79.92% swabs analysed thus far and it is concluded that Bd seems absent in the study region, a result which has massive conservation implications for the region. v Despite the fact that the Okavango Delta has benefitted from conservation and tourism efforts in the past, the system and its biodiversity remains threatened and effective conservation management strategies must be devised and implemented to ensure its preservation. Key words: Amphibians; Diversity; Breeding; Chytrid; Okavango Delta. Xigera Lagoon, the Okavango Delta vi TABLE OF CONTENTS TITEL PAGE i PREFACE AND ACKNOWLEDGEMENTS ii ABBREVIATIONS USED IN THE TEXT iv EXPLANATION OF NON-ENGLISH TERMS iv ABSTRACT v TABLE OF CONTENTS vii CHAPTER ONE: GENERAL INTRODUCTION 1.1 The Okavango Delta: a flood-pulsed ecosystem 1 1.2 Study area 4 CHAPTER TWO: AMPHIBIAN SPECIES DIVERSITY IN THE OKAVANGO DELTA: THE EFFECT OF ISOLATION 2.1 INTRODUCTION 2.1.1 Amphibian diversity and conservation: a closer look at 14 the Okavango Delta 2.1.2 The effect of isolation on amphibian diversity 17 2.1.3 Hypothesis and objectives 19 2.2 METHODOLOGY 2.2.1 Assessment of locality isolation 20 2.2.2 Sampling effort 20 2.2.3 Collection of amphibian species in the Okavango Delta 21 2.2.4 Voucher collection and preparation of voucher specimens 24 2.2.5 Assessment of amphibian community composition per locality 25 2.2.6 Statistical evaluation of differences in community composition 27 2.2.7 Investigation of species occurrence related to habitat type 28 2.2.8 Investigation of habitat types available at each locality 29 2.3 RESULTS 2.3.1 Degree of locality isolation 30 vii 2.3.2 Species richness for the Okavango Delta: actual versus 32 expected 2.3.3 Species richness per locality 40 2.3.4 Species diversity and evenness among localities 43 2.3.5 Community composition 44 2.3.6 Relation between species occurrence and habitat type 45 2.3.7 Physiographic regions and the availability of amphibian 50 habitat types 2.4 DISCUSSION 2.4.1 Amphibian species richness in the Okavango Delta 54 2.4.2 Isolation as a driver for amphibian community composition 55 2.4.3 Habitat availability as a driver for species richness 58 2.4.4 Investigation of available habitat types per locality 60 CHAPTER THREE: HYDROLOGY AS DRIVER FOR AMPHIBIAN BREEDING IN THE OKAVANGO DELTA 3.1 INTRODUCTION 3.1.1 Reproductive patterns of amphibians in Southern Africa 62 3.1.2 The function of anuran vocalisation in the reproductive process 64 3.1.3 Nutrient flow and productivity in the Okavango Delta 66 3.1.4 Hypothesis and objectives 67 3.2 METHODOLOGY 3.2.1 Literature review of breeding seasons for Okavango Delta 68 species 3.2.2 Observation of breeding indicators under natural conditions 68 3.2.3 Assessment of breeding classes and seasons for Okavango 70 Delta species 3.2.4 Comparisons and differences in literature based and present 70 study breeding class composition 3.2.5 Correlations of breeding activity with rainfall and flood pulse 71 3.3 RESULTS 3.3.1 Literature review of breeding seasons for Okavango Delta 72 species 3.3.2 Breeding classes of Okavango Delta species 73 viii 3.3.3 Comparisons
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