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Color and Communication in Habronattus Jumping Spiders

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Color and Communication in Habronattus Jumping Spiders Color and Communication in Habronattus Jumping Spiders: Tests of Sexual and Ecological Selection by Lisa Anne Taylor A Dissertation Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy Approved January 2012 by the Graduate Supervisory Committee: Kevin McGraw, Chair Ronald Rutowski David Clark James Johnson John Alcock ARIZONA STATE UNIVERSITY May 2012 ABSTRACT Differences between males and females can evolve through a variety of mechanisms, including sexual and ecological selection. Because coloration is evolutionarily labile, sexually dichromatic species are good models for understanding the evolution of sex differences. While many jumping spiders exhibit diverse and brilliant coloration, they have been notably absent from such studies. In the genus Habronattus, females are drab and cryptic while males are brilliantly colored, displaying some of these colors to females during elaborate courtship dances. Here I test multiple hypotheses for the control and function of male color. In the field, I found that Habronattus males indiscriminately court any female they encounter (including other species), so I first examined the role that colors play in species recognition. I manipulated male colors in H. pyrrithrix and found that while they are not required for species recognition, the presence of red facial coloration improves courtship success, but only if males are courting in the sun. Because light environment affects transmission of color signals, the multi- colored displays of males may facilitate communication in variable and unpredictable environments. Because these colors can be costly to produce and maintain, they also have the potential to signal reliable information about male quality to potential female mates. I found that both red facial and green leg coloration is condition dependent in H. pyrrithrix and thus has the potential to signal quality. Yet, surprisingly, this variation in male color does not appear to be important to females. i Males of many Habronattus species also exhibit conspicuous markings on the dorsal surface of their abdomens that are not present in females and are oriented away from females during courtship. In the field, I found that these markings are paired with increased leg-waving behavior in a way that resembles the pattern and behavior of wasps; this may provide protection by exploiting the aversions of predators. My data also suggest that different activity levels between the sexes have placed different selection pressures on their dorsal color patterns. Overall, these findings challenge some of the traditional ways that we think about color signaling and provide novel insights into the evolution of animal coloration. ii DEDICATION To my parents, who have always encouraged my curiosity in creepy, crawly things iii ACKNOWLEDGMENTS I would first like to thank my advisor, Kevin McGraw, whose support, guidance, and ideas have provided me with an amazing graduate school experience. The rest of my dissertation committee - John Alcock, David Clark, Chad Johnson, and Ronald Rutowski – has also provided invaluable feedback, ideas, and enthusiasm. I would like to thank all of the members of the McGraw and Rutowski labs. In particular, Melissa Meadows, Mike Butler, and Matthew Toomey have been wonderful lab- and office mates, without whom graduate school would not have been nearly as productive and fulfilling. The rest of the McGraw and Rutowski lab members – Melissa Amarello, Mathieu Giraudeau, Nikos Lessios, Rusty Ligon, Nate Morehouse, Kim Pegram, Brett Seymoure, Melissah Rowe, and Melinda Weaver – have also provided invaluable insight and support over the years. This research would have been impossible without the hard work of numerous undergraduate assistants, including Karrina Domke, Jim Grieco, Joe Kling, Lana Hall, Garrett Lewis, Allison Lopez, Nathan Mills, Sarah Moratto, Nik Moseby, Megan Ponce, Liana Ramirez, Meghan Still, and Claudia Torres. Numerous friends and colleagues have made my time at ASU not only intellectually rewarding, but a lot of fun too. Thanks to Monamie Bhadra, Rebecca Clark, Gustavo Garduno, Jason Hurst, Akito Kawahara, Dani Moore, Jeff Propster, Chris Rader, Emily Schoerning, and the entire SoLS graduate community. iv Funding was provided by a National Science Foundation Graduate Research fellowship and grants from the Animal Behavior Society, the American Arachnological Society, Sigma Xi (national and ASU chapter), the ASU Graduate and Professional Students’ Association, and the ASU Office of Research and Training Initiatives. Finally, thank you to the Rio Salado Habitat Restoration Project and Schnepf Farm for permission to collect spiders and conduct fieldwork on their property. v TABLE OF CONTENTS Page LIST OF TABLES ...................................................................................................... ix LIST OF FIGURES .................................................................................................... xi PREFACE...... ........................................................................................................... xiv CHAPTER 1 MISDIRECTED COURTSHIP IN A NATURAL COMMUNITY OF COLORFUL HABRONATTUS JUMPING SPIDERS .................. 1 Introduction ......................................................................................... 2 Methods ............................................................................................... 7 Results ............................................................................................... 13 Discussion ......................................................................................... 16 2 SEX-DIFFERENCES IN DECEPTION: CAN ACTIVITY PATTERNS OR MICROHABITAT USE EXPLAIN SEXUALLY DIMORPHIC DORSAL COLOR PATTERNS IN HABRONATTUS JUMPING SPIDERS? ..................................... 33 Introduction ....................................................................................... 34 Methods ............................................................................................. 42 Results ............................................................................................... 48 Discussion ......................................................................................... 51 3 ONTOGENETIC COLOR CHANGE IN SEXUALLY DICHROMATIC HABRONATTUS PYRRITHRIX JUMPING SPIDERS ........................................................................................ 69 vi Introduction ....................................................................................... 70 Methods ............................................................................................. 75 Results ............................................................................................... 81 Discussion ......................................................................................... 82 4 MALE ORNAMENTAL COLORATION IMPROVES COURTSHIP SUCCESS IN A JUMPING SPIDER, BUT ONLY IN THE SUN… ........................................................................................ .... 94 Introduction ....................................................................................... 95 Methods ........................................................................................... 100 Results ............................................................................................. 114 Discussion ....................................................................................... 115 5 NATURAL VARIATION IN CONDITION-DEPENDENT DISPLAY COLORATION DOES NOT PREDICT MALE COURTSHIP SUCCESS IN A JUMPING SPIDER ......................................... 132 Introduction ..................................................................................... 133 Methods ........................................................................................... 136 Results ............................................................................................. 148 Discussion ....................................................................................... 150 6 CONCLUDING REMARKS ............................................................. 163 Sexual selection .............................................................................. 163 Ecological selection ........................................................................ 167 Salticids as models for future work ................................................ 169 REFERENCES ...................................................................................................... 171 vii APPENDIX A ANIMAL COLORATION: SEXY SPIDER SCALES ................ 202 B CONDITION DEPENDENCE OF MALE DISPLAY COLORATION IN A JUMPING SPIDER ............................... 205 C SUPPLEMENTARY MATERIAL FOR CHAPTER 2 ................ 220 D SUPPLEMENTARY MATERIAL FOR CHAPTER 4 ............... 222 E SUPPLEMENTARY MATERIAL FOR CHAPTER 5 ................ 225 F COAUTHOR PERMISSIONS FOR INCLUSION OF PUBLISHED WORKS ....................................................................................... 227 viii LIST OF TABLES Table Page 1. Spearman rank correlations between female habitat preference and the preference of conspecific males in four species of sympatric Habronattus (H. clypeatus, H. hallani, H. hirsutus, and H. pyrrithrix). ............................................................................................................ 31 2. Spearman rank correlations between female habitat preference and the preference of conspecific males in three litter-dwelling species of Habronattus
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