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Biology of Wood-Eating Sea Urchins

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Biology of Wood-Eating Sea Urchins EVOLUTIONARY BIOLOGY OF WOOD-EATING SEA URCHINS (TEMNOPLEURIDEA: TRIGONOCIDARIDAE) A Thesis submitted to the faculty of . , San Francisco State University In partial fulfillment of the requirements for b 'O L - 'heDegree Master of Science In Biology: Ecology, Evolution, and Conservation Biology by Catherine Flelen Brett San Francisco, California January 2017 Copyright by Catherine Helen Brett 2017 CERTIFICATION OF APPROVAL I certify that I have read Evolutionary Biology of Wood-Eating Sea Urchins (Temnopleuridea: Trigonocidaridae) by Catherine Helen Brett, and that in my opinion this work meets the criteria for approving a thesis submitted in partial fulfillment of the requirement for the degree Master of Science in Biology: Ecology, Evolution and Conservation Biology at San Francisco State University. Rich Mooi, Ph.D. Research Professor (SFSU), Curator (CAS) WV <■ Terrence Gosliner, Ph.D. Research Professor (SFSU), Curator (CAS) Greg Spice^yPh.D. Professor of Biology, SFSU EVOLUTIONARY BIOLOGY OF WOOD-EATING SEA URCHINS (TEMNOPLEURIDEA: TRIGONOCIDARIDAE) Catherine Helen Brett San Francisco, California 2017 Echinoids comprise a diverse group of Echinoderms composed of around 1000 extant species. The sea urchin family, Trigonocidaridae exemplifies echinoid diversity, being found in almost every marine ecoregion from the intertidal zone to depths over 3,000 m. Some of the Trigonocidaridae are also known wood-eaters, an unusual behavior previously recorded in only three sea urchin species. The family is also characterized by miniature body size and sculpturing of the test. This study is the first phylogeny of the Trigonocidaridae inferred from morphological and molecular evidence. All morphological analyses support the monophyly of the family Trigonocidaridae as well as the monophyly of the clades Trigonocidaris + Genocidaris + Desmechinus, and Hypsiechinus + Prionechinus + Asterechinus (which is monotypic). The molecular analysis does not contradict the morphological analyses and shows strong support for a relationship between Asterechinus and Prionechinus. The evolution of wood-eating and the biogeography of the Trigonocidaridae are explored using this phylogeny. I certify that the Abstract is a correct representation of the content of this thesis. Chair, Thesis Committee Date ACKNOWLEDGEMENTS I would like to thank Rich Mooi for enthusiastically accepting me into his lab and for his thoughtful guidance and encouragement. I would also like to thank the other members of my committee, Terry Gosliner and Greg Spicer for their support. This project was financially supported by the Graduate Student Council of Biology at SFSU. I thank Adam Baldinger at the Harvard Museum of Comparative Zoology, Marc Eleaume at the Paris Museum of Natural History, and Tom Schiotte at the Zoological Museum of the University of Copenhagen for specimen loans, without which this project would not be possible. I am grateful to Anna Sellas who spent hours training me and answering all of my questions, and to Boni Cruz and Claudia Rocha who generously gave their time to train and mentor me in the Center for Comparative Genomics. I thank my lab-mates, Kelly Markello and Kristen Vollrath, for their friendship, feedback and guidance, and Chrissy Garcia and Erika Garcia for training me on the Leica microscopes. I extend my gratitude to all my friends for their crisp wit in keeping me light-hearted and laughing. And finally, I would like to acknowledge my family, without whom I could not have completed this project, with special thanks to Matt and Kacey, and Auntie Paula and Uncle Patrick. And for their love and encouragement, I'd like to thank my parents Bob and Roberta Brett. Most of all to my mom, who is my biggest cheerleader and endlessly inspiring, my deepest thanks. v TABLE OF CONTENTS List of Table........................................................................................................................... viii List of Figures...........................................................................................................................ix List of Appendices................................................................................................................... xi Introduction.................................................................................................................................1 Taxonomic History.......................................................................................................3 Distribution................................................................................................................... 3 Important Morphological Traits...................................................................................4 Wood-eating in the Deep Sea...................................................................................... 8 Significance of the Research...................................................................................... 11 Research Objectives.................................................................................................... 12 Materials and Methods............................................................................................................ 14 Morphological Analyses.............................................................................................16 Molecular Analysis..................................................................................................... 17 Phylogenetic Analysis................................................................................................ 20 Gut Content Analysis..................................................................................................22 Character Descriptions and Character State Coding................................................23 Results...................................................................................................................................... 28 Morphological Parsimony Analysis.......................................................................... 28 Reweighted Analysis..................................................................................................29 Maximum Likelihood Molecular Analysis...............................................................30 Phylogenetic Relationships with Trigonocidaridae................................................. 31 Gut Dissection Analysis............................................................................................. 32 Discussion................................................................................................................................ 35 Phylogeny from Morphological Analysis.................................................................36 Molecular Relationships............................................................................................ 39 Biogeography.............................................................................................................. 40 Wood-eating Behavior............................................................................................... 42 Future Research..........................................................................................................45 References................................................................................................................................ 47 Appendices............................................................................................................................... 92 LIST OF TABLES Table Page 1. List of extant species of Trigonocidaridae...................................................... 53 2. List of specimens shown in images..................................................................54 3. List of specimens used for gut content analysis................................. 55 4. List of MNHN voucher specimens for molecular analysis............................56 5. List of NCBI voucher specimens for molecular analysis................................ 57 viii LIST OF FIGURES Figures Page 1. Map of trigonocidarid distribution....................................................................... 58 2. Regular sea urchin anatomy..................................................................................59 3. Detail of Desmechinus versicolor, ZMUC ECH-330.........................................60 4. Side view of Asterechinus elegans, ZMUC ECH-700....................................... 61 5. Detail of Trigonocidaris versicolor, ZMUC ECH-329..................................... 62 6. Detail of Prionechinus forbesianus, NMNH E5684.......................................... 63 7. Detail of Asterechinus elegans, MNHN IE-2013-AT100-4338........................64 8. Asterechinus elegans, MNHN IE-2013-AT 100-4338........................................65 9. Desmechinus anomalus, Holotype, MCZ ECH-4635....................................... 66 10. Desmechinus rufus, Syntype, MCZ ECH-4735.................................................. 67 11. Desmechinus versicolor, Syntype, ZMUC ECH-330.........................................68 12. Genocidaris maculata, MCZ ECH-8015.............................................................69 13. Hypsiechinus coronatus, Syntype, ZMUC ECH-10...........................................70 14. Prionechinus agassizii, MCZ ECH-3935............................................................71 15. Prionechinus forbesianus, Syntype, MCZ ECH-3941.......................................72 16. Prionechinus sagittiger, ZMUC ECH-715..........................................................73
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