The Behavioral Ecology and Territoriality of the Owl Limpet, Lottia Gigantea
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THE BEHAVIORAL ECOLOGY AND TERRITORIALITY OF THE OWL LIMPET, LOTTIA GIGANTEA by STEPHANIE LYNN SCHROEDER A DISSERTATION Presented to the Department of Biology and the Graduate School of the University of Oregon in partial fulfillment of the requirements for the degree of Doctor of Philosophy March 2011 DISSERTATION APPROVAL PAGE Student: Stephanie Lynn Schroeder Title: The Behavioral Ecology and Territoriality of the Owl Limpet, Lottia gigantea This dissertation has been accepted and approved in partial fulfillment of the requirements for the Doctor of Philosophy degree in the Department of Biology by: Barbara (“Bitty”) Roy Chairperson Alan Shanks Advisor Craig Young Member Mark Hixon Member Frances White Outside Member and Richard Linton Vice President for Research and Graduate Studies/Dean of the Graduate School Original approval signatures are on file with the University of Oregon Graduate School. Degree awarded March 2011 ii © 2011 Stephanie Lynn Schroeder iii DISSERTATION ABSTRACT Stephanie Lynn Schroeder Doctor of Philosophy Department of Biology March 2011 Title: The Behavioral Ecology and Territoriality of the Owl Limpet, Lottia gigantea Approved: _______________________________________________ Dr. Alan Shanks Territoriality, defined as an animal or group of animals defending an area, is thought to have evolved as a means to acquire limited resources such as food, nest sites, or mates. Most studies of territoriality have focused on vertebrates, which have large territories and even larger home ranges. While there are many models used to examine territories and territorial interactions, testing the models is limited by the logistics of working with the typical model organisms, vertebrates, and their large territories. An ideal organism for the experimental examination of territoriality would exhibit clear territorial behavior in the field and laboratory, would be easy to maintain in the laboratory, defend a small territory, and have movements and social interactions that were easily followed. Lottia gigantea , the owl limpet, is just such a model animal. With a small territory (< 900 cm 2) and slow movements (3 mm/min), the interactions of several L. gigantea can be continuously and simultaneously monitored. Using time-lapse photography, experiments were conducted to observe behaviors of L. gigantea , ranging from how L. gigantea form home ranges to how territorial L. gigantea interact. Lottia gigantea formed home ranges within four weeks, returning to a iv home scar after each foraging cycle. To determine whether L. gigantea returned to areas with greater food resources, three different algal density treatments were used, and individuals were monitored to see which tiles they frequented the most. Lottia gigantea actually avoided areas with a thick algal covering, potentially due to the loss of suction they experienced while moving across algae. When L. gigantea established territories, home ranges overlapped considerably. Two individuals were placed in one arena, under the assumption that a dominance hierarchy would be established. Dominant status was predetermined, and in four of the seven dyads both individuals were evasive. When subjected to territorial encounters for two weeks, L. gigantea avoided areas where they experienced agonistic losses. Mucus may serve as an olfactory cue to define territorial boundaries. Individuals avoided tiles with conspecific’s mucus more often than tiles with self-mucus or no mucus. v CURRICULUM VITAE NAME OF AUTHOR: Stephanie Lynn Schroeder PLACE OF BIRTH: Green Bay, Wisconsin DATE OF BIRTH: December 26, 1978 GRADUATE AND UNDERGRADUATE SCHOOLS ATTENDED: University of Oregon, Eugene University of Wisconsin, Milwaukee DEGREES AWARDED: Doctor of Philosophy, Biology, 2011,University of Oregon Bachelor of Science, Biology, 2001, University of Wisconsin, Milwaukee AREAS OF SPECIAL INTEREST: Behavioral Ecology Marine Biology Invertebrate Biology PROFESSIONAL EXPERIENCE: Research Assistant, University of Oregon, 2009-2011 Supervisor, Alan Shanks Teaching Assistant, University of Oregon, 2005-2009 Supervisors: Richard Emlet, Alan Shanks, Elizabeth Whitchurch, Sandra Brooks, Cynthia Trowbridge NSF GK-12 Fellow, Oregon Institute of Marine Biology, 2005-2007 Supervisors: Jan Hodder, Patricia Mace, Alan Shanks Research Assistant, University of California, Davis, 2001-2003 Supervisor, Susan Williams vi GRANTS, AWARDS, AND HONORS: Student Scholarship, Green Acres Grange, 2010 Travel Grant, Malacological Society of London, 2009 Research Grant, Pacific Northwest Shell Club, 2009 Phyllis Para Talus Presidential Fellowship, University of Oregon, 2009 Research Grant, Conchologists of America, 2008 Research Grant, Western Society of Malacologists, 2007 Research Grant, Dr Earl H. Myers and Ethel M. Myers Oceanographic and Marine Biology Trust 2007 Student Scholarship, USA Funds, 2007 Neil Richmond Fellowship, Oregon Institute of Marine Biology, 2007 Graduate Research Fellowship Honorable Mention, National Science Foundation, 2006 Research Grant, Sigma Xi Grants in Aid of Research, 2006 PUBLICATIONS: Schroeder SL, LaFerriere A, Ward J et al. (2009) Beach Hoppers: Inquiry-based learning while having Fun! Clearing magazine, Compendium 2009:Environmental Education in the Pacific Northwest Pgs 20-22 Williams SL and Schroeder SL (2004) Eradication of the invasive seaweed Caulerpa taxifolia by chlorine bleach. Marine Ecology Progress Series 272:69-72 vii ACKNOWLEDGMENTS First and foremost, I would like to thank my advisor, Alan Shanks. Alan always had time to listen to questions and concerns about my project. In addition, his ability to think outside the box pushed me to be a creative scientist. Second, I would like to thank my dissertation committee, Mark Hixon, Bitty Roy, Craig Young and Frances White, for their suggestions, advice and support throughout the years. Special thanks goes to Frances White who helped me tremendously with my statistical analyses and set aside time when I would frantically call her saying I needed help. Bill Wright was an enormous help whenever I had limpet questions or needed a sounding board, in addition to supplying me with limpets. Many thanks to the OIMB staff for keeping things running smoothly at the lab. Jan Hodder and Trish Mace did a fantastic job with the NSF GK-12 program at OIMB and their encouragement made me realize I want to continue doing educational outreach and informal science education. Barb Butler always managed to locate obscure references for me and kept me motivated as my informal dissertation coach. She only made me cry once. Of course I would never be where I am without the love and support of my family. My mom and dad encouraged me since I was a child to do whatever I wanted and were never bothered that I wanted to play in the mud and collect insects rather than play with dolls. Jeremy and Katie, my brother and sister-in-law, have been so supportive and I cannot express in words the gratitude I feel. And a special thanks to my amazing friends, both near and far for always being supportive and cheering me up. Grand ladies of the trifecta, you kept me sane and I love viii you. The thought of leaving my close-knit community reduces me to tears, but I know we will be forever bonded. And a huge thanks goes to the organizations that supported my research: the Malacological Society of London, the Pacific Northwest Shell Club, the University of Oregon, the Conchologists of America, the Green Acres Grange, the Western Society of Malacologists, the Dr. Earl H. Myers and Ethel M. Myers Oceanographic and Marine Biology Trust, USA Funds, the Neil Richmond Fellowship, Sigma Xi and the National Science Foundation under grant no. 0338153. ix This dissertation is dedicated to Dave Larson. Dave was one of my favorite uncles. He would always give me a hard time about becoming one of the ivory tower elite, despite being a teacher himself. This gentle ribbing kept me grounded and humble throughout graduate school. Dave passed away from ALS (Lou Gehrig’s disease) last year. He fought a good fight and his memory lives on. x TABLE OF CONTENTS Chapter Page I. GENERAL INTRODUCTION ................................................................................ 1 Gardening............................................................................................................... 1 Homing .................................................................................................................. 2 Territorial Behavior in the Field ............................................................................ 2 Lottia gigantea Territorial Behavior..................................................................... 3 Scope and Objectives of This Dissertation............................................................ 5 II. METHODS FOR LONG-TERM MAINTENANCE AND MONITORING OF THE OWL LIMPET, LOTTIA GIGANTEA ............................................................ 8 Introduction............................................................................................................ 8 Materials and Methods........................................................................................... 11 Animal Husbandry........................................................................................... 11 Time-Lapse Photography................................................................................. 13 Results.................................................................................................................... 14 Discussion.............................................................................................................