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Life History and Physiologocal Ecology of the Lizard Cordylus

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Life History and Physiologocal Ecology of the Lizard Cordylus II I - LIFE HISTORY AND PHYSIOLOGICAL ECOLOGY OF THE LIZARD, CORDYLUS GIGANTEUS. BY I JOHANNES H. VAN WYK Thesis submitted in fulfilment of the requirements for the degree of DOCTOR OF PHILOSOPHY University of Cape Town in the Department of Zoology, Faculty of Science, University of Cape Town. JUNE 1992 The copyright of this thesis vests in the author. No quotation from it or information derived from it is to be published without full acknowledgement of the source. The thesis is to be used for private study or non- commercial research purposes only. Published by the University of Cape Town (UCT) in terms of the non-exclusive license granted to UCT by the author. University of Cape Town l To my late father, Professor J. H. van Wyk, who in his short life could only dream about me walking the road of a scientist, the wonderful example he set will always be a great inspiration to me, and to my late my late parents-in-law, J. P. and J. F. Franken who patiently waited, but never lived to see also one of their dreams come true. University of Cape Town ii DECLARATION The research reported in this thesis is original work, completed by me between 1984 and '( 1990 under supervision of the University of Cape to.wn. The pata collection, analysis and interpretations are my own, and any assistance I received is fully acknowledged. University of Cape Town iii ACKNOWLEDGEMENTS I gratefully acknowledge the many people and institutions who supported me during this study. Professor Gideon Louw, for accepting me as a student in the _first place, and for all the encouragement and supervision he gave me during the fieldwork stages of this project. Dr Scott Turner, for resuming the supervision of my thesis following the departure of Prof. Gideon Louw. Dr Sue Nicolson, my third supervisor, for valuable advice and encouragement while writing my thesis, but most of all, for patiently and without any delay reading my thesis. I am greatly indebted to her for improving the literary presentation of my writings. My external examiners, Prof L. J. Guillette (Florida University, USA), Prof R. Shine (University of Sydney, Australia) and Prof A. Bauer (Villanova University, USA) for valuable suggestions. The Council and Directors, Drs C. M. Engelbrecht and C. D. Lynch of the National Museum for encouragement and support during the years of my employment. I greatly appreciate the ample opportunity for field research, the use of museum facilities, most of the financial support for field studies, and later permission to use the study collection. The Orange Free State UniversityDepartment of Nature of Conservation, Cape Townespecially Dr Lucas Stolz, for advice, support and permission to conduct this project. Mr C.J. Retief of the farm Greenlands for allowing me to collect lizards on his farm. Mr Wynand Louw and his family for their hospitality, continued interest and all their support while conducting research on their farms. Wynand is the perfect host for a field researcher, who, at all times without hesitation, provided valuable logistic support. iv Sincere thanks are owed to my research assistants at the National Museum, Trudie Saaiman, Rod Douglas, Mike Bates, Frans and Simon who provided valuable field and laboratory assistance during this study. , I have great appreciation for their dedication and all the sacrifices they made during the course of this project. Department of Chemical Pathology, University of the Orange Free State in particular Professor W.J.H. Vermaak for kindly providing facilities, in the use of the radio­ immuno-assay laboratory and the SMAC analytic system. Mr T. Muller for his invaluable assistance during the hormone assays. Department of Soil Science, University of the Orange Free State, in particular Prof. Burger for the use of their laboratory facilities. Department of Geology, University of the Orange Free State, m particular, Prof Nie Grobbelaar, for the use of their surveying field equipment. Public Works Department for the use of scafholding to build an observation tower in the field. Department of Zoology, University of Stellenbosch for allowing me to finalize this study. The South African Weather Bureau for kindly providing climatic data of the study areas. I am also indepted to the following people: - Dr Schalk Louw and his staff in the Entomology Section at the National Museum for helping with the identification of insects. - Prof S. Maritz forUniversity all the discussions of on Cape growth models Town and the mathematical development of an oscillating growth model. - Mandy Lombard for spectrophotometric measurements of lizard skins. - Dirk Versveld for being so helpful with the MCS datalogger and tape reader. - Mr B.P. Ludick for valluable information on the agriculture of the High Veld Region. - Johan Watson for many discussion during the planning of this project and also for field assistance when I needed it most. v -,All the staff that worked at the National Museum during my employment for the invaluable assistance and support they gave me. A special word of thanks to the librarians, late Tharina Koole and Millie Bronkhorst for all their help. - LeFras Mouton for the use of his printer and for all the times he took responsibility for my duties so that I could work on my thesis. - Mr A. Lintveld for all his help since my return to Stellenbosch. - Marietjie Steyn (help with art work and diagrams), Shirley de Viliers (help with typing and printing) and Mrs. J. Botha for all the help in the Zoology library at Stellenbosch. - Dr F. Ellis who helped me understand the pedology terminology. A special word of thanks also to my dear mother and sister as well as all my relatives-in-law who always supported and encouraged me. Finally, my most special thanks to my wife, Irna and two sons Jos and Franken, for all their patience, sacrifices and perpetual encouragement all these years. University of Cape Town vi ABSTRACT LIFE HISTORY AND PHYSIOLOGICAL ECOLOGY OF THE LIZARD, CORDYLUS GJGANTEUS. J.H. van Wyk, Zoology Department, University of Cape Town, Rondebosch 7700, South Africa. 1992. Cordylus giganteus is a large, terrestrial, viviparous lizard, endemic to the Highveld grasslands of South Africa. Its distribution is limited and its conservation status is vulnerable. Autopsy and mark-recapture methods were used to study the seasonal aspects of its reproductive cycle, diet, energy reserves, growth, population dynamics, daily activity and thermoregulation. Reproduction is distinctly seasonal in both sexes. Females may reproduce biennially. Vitellogenesis commenced in autumn (March), and continued through hibernation with ovulation in spring (October). Two or three young are born in autumn. A functional placenta is implicated. Seasonal steroid hormone profiles are presented. Males exhibit a postnuptial spermatogenetic cycle. Spermatogenesis commences in spring with peak spermiogenesis in autumn and testicular regression following in late autumn. Spermatozoa are stored in the epididymis and ductus def erens for seven to eight months. A bimodal plasma testosterone profile is reported, consistent with spermiogenesis in autumn and mating behaviour in spring. C. giganteus feeds during 8 months of the year and prefer Coleoptera as prey. Fat bodies are utilized for winter maintenance and reproduction. Hatchlings grow 20-30mm during the first year and maximum growth rates Universityoccur in summer. Males of andCape females Townattain sexual maturity at about 165 mm SVL in the fourth year. Seasonal affects on growth rate resulted in poor fit by either logistic-by-length or von Bertalanffy models and a seasonal oscillating model was introduced. Adult males are smaller than females; head sizes are the same but allometric slopes differed significantly. Population size and structure remained stable in the study area. Densities ranged from 9 - 11 lizards/ha. The age structure is marked by the low relative abundance of juveniles. Survivorship during the first year varied among years. Mortality was highest during summer months rather than winter months. Average annual survival of adults 'was high, but varied with sex and years (ranged from 58%-80% ). A life table yielded a net reproductive rate (Ro= 1) sufficient to sustain the population, if the vii reproductive life of an adult female is at least 12 years. Lizards remain in their burrows during winter. In summer, activity was bimodal on sunshine days but unimodal on cool overcast days. Body temperature is regulated by behavioural means (postural and orientation changes) and by shuttling to the cool burrow microclimate. The life history strategy corresponds partially to that of K-selection. University of Cape Town viii CONTENTS CHAPTER 1: GENERAL INTRODUCTION 1.1. LITERATURE REVIEW 1 1.1.1. Discovery, Nomenclature and General Description 1 1.1.2. Distribution and General Conservation Status 3 1.1.3. Existing Conservation Measures 11 1.1.4. Recent Research 11 1.2. 0IUECTIVES OF THE STUDY 13 CHAPTER 2: STUDY AREAS, HABITAT AND GENERAL PROCEDURES 2.1. LOCATION 15 2.2. PHYSIOGRAPHY 15 2.2.l. Geology 15 2.2.2. Pedology 16 2.2.3. Climate 16 2.2.3.1. Highveld Region 16 2.2.3.2. Harrismith 18 2.2.3.3. Bethlehem 20 2.3. VEGETATION 25 2.4. LIZARD BURROWSUniversity of Cape Town 25 2.5. LIZARD Cou:ECTION 33 2.5.l. Field Collecting and Transportation 33 2.5.2. Autopsy and Storage 33 2.6. MARK-RECAPTURE STUDY 35 ix CHAPTER 3: FEMALE REPRODUCTION: TIMING OF THE REPRODUCTIVE CYCLE 3.1. INTRODUCTION 36 3.2. MATERIAL AND METHODS 38 3.2.1. Study Area and Lizards 38 3.2.l.l. Autopsy and Measurement 39 3.2.1.2. Mark - Recapture Study 40 3.2.2. · Environmental Correlates and Statistics. 40 3.3. RESULTS 41 3.3.1. Size at Sexual Maturity 41 3.3.2. Reproductive Condition 41 3.3.3.
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