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31295005656821.Pdf (1.518Mb) FORAGING ECOLOGY OF A NEOTROPICAL FOLIVORE, LAMPONIUS PORTORICENSIS REHN (PHASMATODEA:PHASMATIDAE) by ELIZABETH SANDLIN SMITH, B.S. A THESIS IN BIOLOGY Submitted to the Graduate Faculty of Texas Tech University in Partial Fulfillment of the Requirements for the Degree of MASTER OF SCIENCE Approved Accepted December, 1989 T3 I tffCj }J4' jlj../, c"/' :J ACKNOWLEOOEMENTS My gratitude extends in many directions and to many special friends. Foremost, I thank Dr. Michael R. Willig for his support, guidance, patience, and nurturing hand throughout my graduate training. I also thank Dr. John C. Zak and Dr. M. Kent Rylander for their encouragement and input in the development and implementation of my research. Next, I wish to thank the Department of Energy and Oak Ridge Associated Universities for awarding me a summer Research Fellowship, and I am grateful to Dr. Robert B. Waide of the Center for Energy and Environment Research in Puerto Rico for his financial assistance and provision for use of equipment and facilities at El Verde. Additionally, I wish to acknowledge and thank Sr. Alejo Estrada for his assistance in establishing my laboratory and work area and also for lending his exhaustive knowledge of the flora and fauna within the Tabonuco Rainforest. My deep thanks for friendship, advice, ideas, and assistance in the field and elsewhere go to Deborah J. Kyrouac, Michael R. Gannon, Gerardo R. Camilo, David R. Ficklen, Robert Huber, Moira J. van Staaden, and Javier Alvarez. I am indebted to my parents for their tireless support and their avid interest in my education. Finally, and most importantly, I wish to express my heartfelt gratitude to my husband, Hunt, who unselfishly and patiently provided the support I needed to see this project to fruition. 11 TABLE OF CONTENTS ACKNOWLEDGEMENTS..................................................................................... n ABSTRACT............................................................................................................. v LIST OFTABLES ................................................................................................... vii LIST OF FIGURES ................................................................................................. viii CHAPTER I. INTRODUCTION AND PRESENTATION OF THE PROBLEMS......................................................................................... 1 Optimal Foraging Theory .......... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .... .. .. .. .. .. .... .... .... .. .. .. .... .. 1 Modifications of Classical OFT....................................................... 2 Herbivores........................................................................................ 2 Factors Affecting Food Choice.................................................................... 3 Toxins and Nutrients........................................................................ 3 Position............................................................................................ 4 Prior Experience............................................................................... 4 Age ................................................................................................... 4 Phasmatids .. .. .. ... .. .. .. .. .. .. .. .. .. .. .. ...... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .... .. .. .. .. .. .. .. .. .. 5 The Problems . .... .. .. .... .. .. .. .... .... .. .. .. .. .. .. ...... .. ...... .. .. .. .. .. .... .. .. .. .. .... .... .. .. .. ...... 6 CHAPTER II. MATERIALS AND METHODS..................................................... 9 Study Site..................................................................................................... 9 Collection and Maintenance ........................................................................ 9 Experimental Plants ......................................................................... 10 Leaf Selection . .. .. .. .. .. .. .. .. .... .. .. .. .... .... .. .... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 11 111 Feeding Trials . .. .. .. .... .. .. .. .. .. ...... .. .. ........ .. .. .. .. .. .. .... .. .. .. .. .. .. ...... ... .. ........ .. .. .. 11 Age- and Sex-Specific Variation ..................................................... 12 Variation Due to Preexposure...................... .. .... .. .. ...... .......... .. .. ...... 13 Intraspecific Food Variation ............................................................ 13 Data Analyses .............................................................................................. 14 CHAPTER III. RESULTS ...................................................................................... 18 Effects of Age- and Sex-Specific Variation ................................................ 18 Effects of Preexposure to One Food............................................................ 19 Effects of Intraspecific Variation in Food Quality ....................................... 21 CHAPTER IV. DISCUSSION................................................................................ 31 Herbivory..................................................................................................... 31 Lamponius portoricensis .............................................................................. 32 Special Problems Faced By Females ........................................................... 33 Implications of Observed Patterns of Consumption ..................................... 34 Influence of Age and Sex on Diet Composition .............................. 34 Influence of Previous Experience on Diet Composition ................. 35 Food Quality as it Relates to Succession ......................................... 35 Effects of Food Variation Within a Forage Species ........................ 37 Monophagy, Polyphagy, and Plant Successional Status. ................. 38 For the Future............................................................................................... 40 LITERATURE CITED ............................................................................................ 42 APPENDIX .............................................................................................................. 47 IV ABSTRACT Recent attention in ecology has focused on responses of consumers to a variety of factors which influence foraging behavior. Herein, I evaluate the responses to different choices of food plants exhibited by an herbivore which is abundant within the Tabonuco Rainforest of Puerto Rico. Previous work indicates that the walking stick, Lamponius portoricensis. appears to forage on a limited array of plant species and selects habitats which contain high densities of Piper treleaseanum. However, in food choice experiments,£,. treleaseanum is its least preferred food. In an attempt to delineate factors contributing to this apparent dilemma, I designed three separate experiments to evaluate ( 1) if walking sticks of different ages or of different sex have different food preferences, (2) if previous exposure to only one food type affects subsequent diet composition, and (3) if walking sticks distinguish between leaves of varying quality from the same plant . Four plants known to be forage for this insect (Dendropanax arboreus, Piper hispidum, £,. treleaseanum, and~ baccifera) were used in food choice experiments. Multivariate analyses reveal that, at different ages, males and females exhibit different patterns of consumption. Likewise, preexposure to only one food influences subsequent diet differently depending upon preexposure regime and sex. In addition, preferences are shown for different qualities of leaves within single forage species. In particular, lower leaves of P. treleaseanum are preferred, while leaves of D. arboreus and .U.. baccifera are eaten indiscriminately. These results corroborate an earlier suggestion that walking sticks v choose their diets to reflect nutritional constraints. Additionally, walking sticks distinguish between plant species, and varying leaf quality within a single species, and also modify their diet to reflect past experience. Thus, they may have an impact upon successional processes and, ultimately, the composition of plant communities within forest light gaps. Vl LIST OF TABLES 1. Annotated results for averaged univariate and multivariate repeated measures analysis of variance of percent consumption data from an experiment showing age-specific variation in food choice ................................................................... 22 2. Annotated results for averaged univariate and multivariate repeated measures analysis of variance of percent consumption data from an experiment showing effects of preexposure on food choice .................................................................. 27 3. Annotated results for averaged univariate and multivariate repeated measures analysis of variance of percent consumption data from an experiment showing variation in intraspecific food choice ................................................................... 28 Vll LIST OF FIGURES 1. Schematic representations of the physiognomy of plant taxa used during feeding trials......................................................................................................... 17 " Histograms showing average percent consumption by L. portoricensis of four foods offered during feeding trials for each age class and sex ............................. 24 3. Histograms showing average percent consumption by L. portoricensis of four foods offered during feeding trials
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