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SHELTER USE by CALCINUS V E W L I , BERMUDA's EX?)Elflc SHELTER USE BY CALCINUS VEWLI, BERMUDA'S EX?)ELflC HELMIT CL4B Lisa Jacqueline Rodrigues A thesis submitted in conformity with the requirements for the degree of Master of Science Graduate Department of Zoology University of Toronto 0 Copyright by Lisa Jacqueline Rodrigues 2000 National Library Bibliothèque nationale du Canada Acquisitions and Acquisitions et Bibliographic Services services bibliographiques 395 Wellington Street 395, rue Wellington Ottawa ON K1A ON4 Ottawa ON K1A ON4 Canada Canada Your irrS Votre mféretut? Our üb Notre rdfénme The author has granted a non- L'auteur a accordé une licence non exclusive licence allowing the exclusive permettant a la National Library of Canada to Bibliothèque nationale du Canada de reproduce, loan, distribute or sell reproduire, prêter, distribuer ou copies of this thesis in microform, vendre des copies de cette thèse sous paper or electronic formats. la forme de microfiche/nlm, de reproduction sur papier ou sur format électronique. The author retains ownership of the L'auteur conserve la propriété du copyright in this thesis. Neither the droit d'auteur qui protège cette thèse. thesis nor substantial extracts bom it Ni la thèse ni des extraits substantiels may be printed or otherwise de celle-ci ne doivent être imprimés reproduced without the author's ou autrement reproduits sans son permission. autorisation. Shelter use by Calcinus vemlli, Bermuda's endemic hennit crab. Master of Science, 2000 Lisa Jacqueline Rodngues Department of Zoology University of Toronto Calcinus vemlli, a hennit crab endemic to Bermuda, is unusual in that it inbabits both gastropod shells (Centhium Iitteratum) and gastropod tubes (Dendropoma irremlare and Dendropoma annulatus; Vermicularia knomi and Vermicularia spirata). Field surveys found significantly more maies inhabiting shells and significantly more females inhabiting tubes. In the laboratory no significant difference was show in shelter choice for males or females. In a second choice experiment, males showed no discrimination between gastropod extract types, and females showed discrimination that related to the shelter they were collected in. In a cornpetition between the sexes, the winner of the intact shell was found to depend upon relative shield length and relative major chela length. Overall, males were found to be better exploitative cornpetitors than fernales. A significant difference in uropod asymmetry was found; tube dwellers were more syrnmetrical than shell dwellers. A shelter manipulation experiment found shelter type to be important in determining asymmetry. The distribution of c. vemlli between shells and tubes may be an example of "making the best of a bad situation," if Polydora sp. is detrimental to the reproductive success of the hennît crab. Although there is jus my name attached to this thesis, there have been many people who have contributed and helped in the past two years. First and foremost, thanks go to both of my supervisors, Dr. David Dunham and Dr. Kathryn Coates. Wztlier I was in Semuda or Toronto, bcth had open doon or were available for on-line discussions. They supplied me with inspiration, as well as advice, and their questions always kept me on my toes. Much of the field work and expenments that 1 conducted during the summer of 1999 could not have been accomplished without Matthew Hammond. As a Bermuda Program student at BBSR Matt helped me with al1 aspects of the project, from collecting and shucking hennit crabs to "sacrificing" gastropods. My work involved transporting animals between Bermuda and Toronto. This was made possible by the issuing of nurnerous irnpon and export permits thanks to Brian Luckhurst and Norbert Simmons of the Department of Fisheries, Coney Island, Bermuda and Mike Souza of BBSR. Mike Souza also measured and cut the Plexiglas pieces for the Y-mare apparatus, and constmaed some glas aquaria. I am also grateful to al1 of the staff of BBSR. Patricia Pocklington introduced me to the wonderful world of polychaetes. She identified the polydorid that 1 found inhabiting the Cerithium litteraîum shells with Calcinus vemlli. Thanks to Dr. Robbie Smith for the use of the research boats, Mako and &a Dance, to Nancy Stevens for the use of Mussel Pie. The boats were captained on most occasions by my dive buddies Sarnantha dePutron and Graham Webster, who were arnazingly able to find just the reef we wanted without the GPS. 1 am especially grateful to Sam for allowing me to search her sites for C. vemlli. Other dive buddies that made SCUBA diving possible were Anne Savage, HaleIy Outerbridge, Eli Woolery, Alex Chequer, and Dr. Joanna Pin. Quincy Burgess, Kevin ?, and Dr. Roger Pocklington had the job of'boat watcher' on nünerous occasions, which involved basking in the Bermuda sun for endless hours on a swaying boat to eventuaily drag aboard exhausted divers and their heavy buckets. In the early months of this Master's program, 1 received incredible support from many people at the Bermuda Aquarium Museum and Zoo (BAMZ). Dr. Annie Glasspool, Billy Mitchell, and Chris Flook helped me collect everything from snapping shrimp to decorator crabs. It was finally Dr. Wolfgang Sterrer who suggested that 1 take a look at C. verrilli; he thought they may result in something interesting, and he was right. 1 received moral support and well wishes from al1 of the stafFat BAMZ. In Toronto, 1 was fortunate to work with two outstanding scientias, Nasreen Rahman and Jan Richards. Erika Crosse helped me make slides and photographie prints of the hermit crabs. The hermit crabs were looked after by several work-audy students: Michelle Herzog, Jennifer Tran, David Punzalan, Mary Sunderland, and Lesia Murray. My roornmates, Deirdre, Denise, and Kathryn patientiy listened to al1 of my hermit crab tales. Finally, 1 thank my parents and brother for their love and support. They have helped me in more ways than 1 can possibly list. This project was funded by a Bermuda Governrnent Scholarship, the Munson Foundation through BBSR, Graduate htern Program of BBSR and a Natural Sciences and Engineering Research Council of Canada research grant to D. Dunham Two CO-authoredscientific papers have been denved f?om some of the material in this thesis: Rodrigues LJ, DW Dunham, and KA Coates (2000) Shelter preferences in the endemic Bermudian hermit crab Calcinus verrilli (Rathbun, 190 1). Cmstaceana 73 : 737- 750. Rodrigues U,DW Dunham, and KA Coates (Submitted) Gastropod shells or gastropod tubes? Shelter choice in the hermit crab Calcinus vemlli. Modem A~oroachesto the Study of Crustacea. The first is paper based on Chapters 1 and 2 of the thesis; the second is based on Chapter 2. 1 designed and conducted ail of the field work and experiments, analyzed the data, did the appropriate library research, and wrote the research papers. Guidance and advice were provided during dl stages by Dr. D.W. Dunham and Dr. K.A. Coates. Title Page -- I Abstract ii ... Acknowledgements - -.-- .-.-.-- ..+-- 111 .. List of Publications --.- -.------.- Table of Contents vi List of Tables - ix List of Figures .-...-.--. xi General Introduction Hennit Crabs: Unique Members of the Subphylum Cmstacea --.....- -.-- 1 Shell Occupancy: Pros and Cons ....- 5 ShelIs: A Lirniting Resource - 7 Corn petition and S hell Exchange ------.-.------.------- 8 Alternative SheIters -- ..-..-...-....--.-- 9 Bermuda's Endemic Hermit Crab ..., ...----.-.-.----.-.- 14 Chapter 1 The distribution of Calcinus vemlli in Bermuda 17 Introduction 17 Methods 21 Updated Distribution - 22 Discussion -29 10. Chapter 2 Shelter preferences in the endemic Bermudian hennit crab, Calcinus verrilli-+--- 3 1 Introduction 3 1 Methods. ------- 34 Intraspecific competition in the endemic Bermudian hermit crab, Calcinus verrilli 61 Resu lts _.........._ ..-....._...-.--.-... 67 75 Discussion .___....-.-..-~-_---.-....-...-.--.- ...........,. .. .. .. 12. Chapter 4 The effects of shelter type on uropod symmetry in Calcinus vemlii 78 Introduction --..78 Met hods. .-- 8 1 13. General Discussion Field Surveys --- 96 Preference for Shelters . 97 Cornpetition for Shelters 99 A Polydorid Anecting Shelter Use 100 Morphological Differences 102 14. Appendix A Regression analyses for gastropod shell measurements and Calcinus vertilli shield length 103 15. Appendix B A new Poivdora sp. (Spionidaz, Polychaeta) found inside the gastrepcd shells of Calcinus O6 vemlli --- -.-. 1 16. References --._ _____I________I____c____._r____..____.._-_____I________I____c____._r____..____..______I________I____c____._r____..____.._...... _____I________I____c____._r____..____.._......_____I________I____c____._r____..____.._ _____I________I____c____._r____..____..______I________I____c____._r____..____.._..._____I________I____c____._r____..____..______I________I____c____._r____..____.._..,.-...-.- 108 viii LIST OF TABLES 1. Hennit crab species known to inhabit shelters alternative to spiraled gastropod shells and the authors who first observed the association Il 2. Sites visited fiom May to September, 1999, and Apd to May, 2000, and the habitats and shelter types occupied by Calcinus vemlli --.-. 24 3. The statisticai significance ofshieid length diffirsnces Semeen Calcinus rerrilli males and females and shelter types, using (a) Mann-Whitney rank sum test and (b) Student's t- test ..-.-- 43 4. The number of males and fernale Calcinus vedli in the three types of shelters coilected in the field - 45 5. Correlation coefficients obtained from linear regressions of Calcinus verrilli shield length for each shell or tube measurernent ....-.-
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