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INTERACTIONS AMONG EGG, LARVAL, and POST-METAMORPHIC PREDATORS of the EAST AFRICAN TREEFROG, Hyperolius Spinigularis

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INTERACTIONS AMONG EGG, LARVAL, and POST-METAMORPHIC PREDATORS of the EAST AFRICAN TREEFROG, Hyperolius Spinigularis SEQUENTIAL PREDATION IN A COMPLEX LIFE-HISTORY: INTERACTIONS AMONG EGG, LARVAL, AND POST-METAMORPHIC PREDATORS OF THE EAST AFRICAN TREEFROG, Hyperolius spinigularis By JAMES RICHARD VONESH A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 2003 Copyright 2003 by James Richard Vonesh To Sophia and Savannah. ACKNOWLEDGMENTS This research could not have been accomplished without the assistance and support of a large number of people. I thank my field assistants, G. Matthews, B. Munizi, S. Mtunguja, A. Kajiru, and S. Balcomb, D. Sutherlin, M. Pauley, and E. Harper for assistance with data collection; C. Sawe, V. Pohjonen of ANR, and K. Howell, C. Msuya, and R. Senzota of the University of Dar es Salaam, for logistical support in the field; V. Clausnitzer for odonate identification; W. Mathis for dipteran identification; and my dissertation committee C. Osenberg (chair), B. Bolker, K. Sieving, L. Chapman, H. Lillywhite, C. St. Mary, and the SOB lab group for valuable feedback though out the entire PhD process, I am a better scientist and person because of our interactions these past years. I thank Tanzanian COSTECH (Permit # 2001-274) and the Division for Forestry and Beekeeping for permission to conduct research and the Tanzania Weather Service, NIMR, Amani, for rainfall data. Financial support was provided by an EPA STAR Fellowship, NSF DDIG DEB-9911965 and the Lincoln Park Zoo. Finally, I thank my family for their encouragement, support, and patience. iv TABLE OF CONTENTS page ACKNOWLEDGMENTS ................................................................................................. iv LIST OF TABLES........................................................................................................... viii LIST OF FIGURES ........................................................................................................... ix ABSTRACT....................................................................................................................... xi CHAPTER 1 INTRODUCTION ........................................................................................................1 2 EGG PREDATION AND PREDATOR-INDUCED HATCHING PLASTICITY IN THE AFRICAN TREEFROG, Hyperolius spinigularis ..............................................8 Introduction...................................................................................................................8 Methods ......................................................................................................................10 Study Site.............................................................................................................10 Hyperolius Abundance, Reproduction, and Egg Survival...................................11 Hatching Plasticity Experiments: Afrixalus fornasini .........................................14 Hatching Plasticity Experiment: Typopsilopa Fly Larvae...................................15 Results.........................................................................................................................16 Hyperolius spinigularis Phenology .....................................................................16 Oviposition Characteristics and Clutch Success .................................................17 Sources of Egg Mortality.....................................................................................19 Egg Predator Effects on Larval Input into the Pond............................................20 Egg Predator Effects on Larval Traits.................................................................21 Discussion...................................................................................................................22 Clutch Success.....................................................................................................22 Egg-stage Predator Effects on Larval-stage Density...........................................24 Egg-stage Predator Effects on Hatching and Larval Traits.................................26 Consequences in Subsequent Life-stages............................................................28 3 CARRY OVER OF PREDATOR EFFECTS ACROSS THREE LIFE-HISTORY STAGES IN AN AFRICAN TREEFROG.................................................................42 Introduction.................................................................................................................42 Methods and Materials ...............................................................................................44 v Site Information...................................................................................................44 Natural History of Hyperolius spinigularis and its Predators .............................44 Experiment 1: Effects of Density on Larval Performance and Postmetamorphic Traits ...................................................................................46 Experiment 2: Size-selective Post-metamorphic Predation ................................48 Results.........................................................................................................................49 Experiment 1: Survival as a Function of Initial Density .....................................49 Experiment 1: Duration of the Larval Stage as a Function of Initial Density.....52 Experiment 1: Size at Metamorphosis as a Function of Initial Density..............54 Experiment 2: Size-specific Metamorph Predation.............................................55 Discussion...................................................................................................................55 4 CONSEQUENCES OF PREDATOR-INDUCED HATCHING PLASTICITY IN AN AFRICAN TREEFROG ......................................................................................66 Introduction.................................................................................................................66 Materials and Methods ...............................................................................................68 Site and System Information ...............................................................................68 Site information............................................................................................68 System information ......................................................................................69 Aquatic larval-stage predators......................................................................70 Consequences of Egg-Stage Predator Effects for Growth and Survival to Metamorphosis.................................................................................................71 Experimental design: density and size/age effects.......................................71 Establishing the egg-predation mediated density effect: N-EP and N+EP1, N+EP2........................................................................................................73 Establishing predator-induced size effect: S-EP and S+EP .............................76 Integrating Patterns of Larval Growth, and Density- and Size-Specific Risk to Predict Larval Mortality...................................................................................77 Larval growth ...............................................................................................77 Effect of density on risk: the functional response ........................................77 Effect of larval size on predation risk ..........................................................79 Combining size and density specific risk.....................................................81 Simulating larval growth and mortality .......................................................82 Results.........................................................................................................................83 Aquatic Predators ................................................................................................83 Larval Survival ....................................................................................................83 Proportion of Surviving Larvae that Reach Metamorphosis...............................84 Mass at Metamorphosis.......................................................................................85 Growth Rates.......................................................................................................86 Density and Size-Specific Risk ...........................................................................86 Comparison of Observed versus Simulated Larval Survival ..............................87 Discussion...................................................................................................................88 vi 5 MULTI-PREDATOR EFFECTS ACROSS LIFE-HISTORY STAGES: NON- ADDITIVITY OF EGG- AND LARVAL-STAGE PREDATION IN AN AFRICAN TREEFROG ...........................................................................................103 Introduction...............................................................................................................103 Materials and Methods .............................................................................................105 Site Information.................................................................................................105 Experimental design ..........................................................................................106 Detecting Multiple Predator Effects..................................................................107
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