INTER- AND INTRASPECIFIC VARIATION IN THE SUPERFAMILY PNEUMOROIDEA Nathan Donelson A Dissertation Submitted to the Graduate College of Bowling Green State University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY December 2007 Committee: Dr. Moira van Staaden, Advisor Dr. Yu Zhou Graduate Faculty Representative Dr. Robert Huber Dr. Karen Root Dr. Dan Wiegmann ii ABSTRACT Dr. Moira J. van Staaden, Advisor This dissertation concerns phenotypic plasticity of male morphology within South African bladder grasshoppers (Orthoptera: Pneumoroidea) from an individual to the species level. Wings and a large inflated abdomen used for acoustic courtship and communication are characteristic of adult male pneumoroids. However, a secondary (‘alternate’) male morph has been identified in three species, which completely lacks the appearance and associated dispersal and signaling behaviors. These characteristics are shared with three monotypic genera formerly raised to accommodate taxa with atypical males. As the aforementioned characteristics are critical for mate localization, the absence of such structures and behaviors demands explanation. The first chapter examines three pneumoroid species that possess both male morphs to determine how similar the uninflated morph is across taxa. The morphology of each species and morph was compared using Multivariate Analysis of Variance (MANOVA) and linear regression. Results show that the morphological differences between inflated and uninflated males is similar across species. This demonstrates that the uninflated phenotype is largely conserved within the Bullacris genus. The second chapter takes a broader comparative approach. It expands the morphometric analysis to investigate three strictly uninflated species and asks whether they are in fact uninflated morphs of inflated species with which they are sympatric. Based on results from chapter 1, we propose that the known differences between morphs can be used to create ‘hypothetical’ uninflated males for any inflated taxa. MANOVA and Discriminant Function iii Analysis (DFA) then compare the hypothetical males to the actual uninflated taxa. Our results suggest that each of the uninflated male taxa can be placed into inflated taxa. This now places dual morphologies in at least three genera and perhaps the entire superfamily. The third chapter uses geographic and morphological data of two widespread species to determine whether environmental characteristics: predict species presence, influence morphology, and form distinct populations. This study also examines whether clinal rules hold across the species. Results from MANOVA, logistic regression, and DFA show that, while environmental conditions are highly predictive of morphology for one species, the second does not display predictable morphological responses to environmental variation. iv I dedicate this to all of my family and friends. I could not have done this without your help. v ACKNOWLEDGMENTS I thank G. Beccaloni, B. Dombrowsky, S. Van Noort, and M. Kieser, curators at the British Museum of Natural History (London), Transvaal Museum (Pretoria), Iziko Museums of South Africa (Cape Town), and the South African National Collection (Rietondale) respectively for access to entomological collections. A. Smith, R. Huber, and S. Anderson provided constructive comments on earlier versions of the manuscript. I thank my graduate committee - M. van Staaden, R. Huber, K. Root, D. Wiegmann, and Y. Zhou - for all of their guidance and advice. Photographer Anthony Bannister supplied valuable locality information concerning his pneumoroid subjects. The South African Department of Environmental Affairs and Tourism and the AGIS provided essential map data. Special thanks to the family of the late Peter Alexander, whose gracious hospitality and logistical support were invaluable to us in South Africa. V. Couldridge and N. Potgeiter for providing key material. Funding for this work was provided in part through a J.P. Scott Center for Neuroscience, Mind & Behavior fellowship, an Oman scholarship, and a National Science Foundation grant (IBN-0091189) to MvS. This work could not have been accomplished without the help of C. Donelson. vi TABLE OF CONTENTS Page GENERAL INTRODUCTION ............................................................................................. 1 The Pneumoroidea (Sensu Flook and Rowell 2000).................................................. 5 CHAPTER I A CROSS-SPECIES COMPARISON OF A MALE POLYPHENISM IN THREE SPECIES OF BLADDER GRASSHOPPERS (ORTHOPTERA: PNEUMOROIDEA) ...... 9 Abstract ..................................................................................................................... 9 Introduction ............................................................................................................... 9 Materials and Methods .............................................................................................. 12 Data Analysis ............................................................................................................ 14 Results ....................................................................................................................... 15 Discussion ................................................................................................................. 17 CHAPTER II USING THE DIFFERENCE BETWEEN MORPHOTYPES IN DUAL-MALE SPECIES TO PREDICT THE MORPHOLOGY OF AN ALTERNATE PHENOTYPE IN SINGLE-MORPH SPECIES ................................................................................................ 21 Abstract ..................................................................................................................... 21 Introduction ............................................................................................................... 22 Materials and Methods .............................................................................................. 26 Data Analysis ............................................................................................................ 27 Results ....................................................................................................................... 29 Discussion ................................................................................................................. 30 CHAPTER III USING MACROECOLOGY TO EXPOSE HOW ENVIRONMENTAL VARIABILITY AFFECTS ORTHOPTERAN MORPHOLOGY ....................................... 35 vii Abstract ..................................................................................................................... 35 Introduction ............................................................................................................... 36 Materials and Methods .............................................................................................. 39 Data Analysis ............................................................................................................ 41 Environmental Predictors of Species Presence ............................................. 41 The Effects of Environmental Variation on Morphology ............................. 42 Determining morphological variation in explicit populations....................... 43 Results ....................................................................................................................... 43 Environmental Predictors of Species Presence ............................................. 43 The Effects of Environmental Variation on Morphology ............................. 44 Determining morphological variation in explicit populations....................... 45 Discussion ................................................................................................................. 47 REFERENCES ..................................................................................................................... 51 APPENDIX A VARIATION IN ADULT LONGEVITY IN A POLYMORPHIC SPECIES .................................................................................................................................... 58 Abstract ..................................................................................................................... 58 Introduction ............................................................................................................... 58 Materials and Methods .............................................................................................. 60 Results ....................................................................................................................... 61 Discussion ................................................................................................................. 62 APPENDIX B TABLES ..................................................................................................... 65 APPENDIX C FIGURES ................................................................................................... 72 viii LIST OF TABLES Table Page 1 Two-way MANOVA of the effects of Body Type and Species on morph .............. 65 2 Mean length values (± standard deviation) for the dual male specimens .................. 66 3 Centroid values for morphological measures in three Bullacris taxa ....................... 67 4 Description of the amount of grasshopper material measured .................................. 68 5 MANOVA test results and the amount of variation explained by each variable ...... 68 6 MANOVA table of environmental effects on body