-UNIVERSITY OF H6WAil LlS~Y EXPLORING NOVEL TAXONOMIC CHARACTER SETS IN THE MOLLUSCA: THE CRIBRARULA CRIBRARIA COMPLEX (GASTROPODA: CYPRAEIDAE) AS A CASE STUDY A DISSERTATION SUBMITTED TO THE GRADUATE DIVISION OF THE UNIVERSITY OF HAWAI'I IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY IN ZOOLOGY MAY 2003 By FABIO MORETZSOHN DE CASTRO JR. Dissertation Committee: E. Alison Kay, Chairperson David W. Greenfield H. Gert de Couet Rebecca Cann Thomas Ramsey © 2003, Fabio Moretzsohn de Castro Jr. 111 ACKNOWLEDGMENTS I am indebted to my academic advisor and mentor, Dr. E. Alison Kay, Emeritus Professor, Department ofZoology, University ofHawaii (UH), for the guidance and inspiration to pursue molluscan taxonomy and evolution. My Ph.D. committee members, E.A. Kay, David W. Greenfield, H. Gert de Couet, Rebecca Cann, and Thomas Ramsey were always very helpful and provided advice in all stages ofmy research. This research was based mostly on museum specimens and would not have been possible without access to several museum collections, and I thank the curators and collection managers who made access to specimens and facilities possible, and for arranging specimen loans: Academy ofNatural Sciences, Philadelphia, PA; the American Museum ofNatural History, New York, NY; the Australian Museum, Sydney, Australia; B. P. Bishop Museum, Honolulu, HI; California Academy of Sciences, San Francisco, CA; Delaware Museum ofNatural History, Wilmington, DE; Field Museum ofNatural History, Chicago, IL; and The National Museum ofNatural History, Smithsonian Institution, Washington, DC. Specimens were also loaned from the following museums, and I thank their staffin arranging the loans: Carnegie Museum ofNatural History, Pittsburgh, PA; Museum National d'Histoire Naturelle, Paris, France; The Natural History Museum, London, United Kingdom; The Natural History Museum ofLos Angeles County, Los Angeles, CA; and The Western Australian Museum, Perth, Australia. IV I am thankful to the many biologists who contributed material: Tom Duda, Curt Fiedler, Lucia Gutierrez, and Dwayne Minton at the University ofHawai'i, and Scott Johnson (Kwajalein), Gustav Paulay (Florida Museum ofNatural History), Cliiudia Baider and Vincent Florens (Mauritius), and Cecilia Osorio (Easter Island, Chile). I am also indebted to many members ofthe Hawaiian Malacological Society generously provided specimens, photographs and encouragement: Don Anderson, Tom and Beatrice Burch, the late C.M. "Pat" Burgess, John Earle (donated a rare specimen ofLyncina leucodon), the late Olive Schoenberg-Dole, Chris Takahashi, Wes Thorsson, Dave Watts, and David Woodman. Financial support from the following institutions was greatly appreciated: Charles Edmondson Research Fund - Grants-in-Aid for Research (UH Dept. Zoology); Conchologists ofAmerica - the Walter Sage award; Delaware Museum ofNatural History - a du PontiMerk visiting fellowship; Graduate Student Organization, UH (travel grant to Species 2000 Workshop, Tsukuba, Japan); Hawaiian Malacological Society student grant; Lerner-Gray Grant for Marine Research (American Museum ofNatural History, New York); Sigma Xi Grants-In-Aid for Research (The Scientific Research Society); and Unitas Malacologica (travel grant to the World Congress ofMalacology, in Washington, DC). I am grateful to the Water Resources Research Center, UH, for supporting me as a Graduate Research Assistant for E. Alison Kay's micromollusk biomonitoring project all these years. Among the many fellow graduate students who made this journey more fun, I remember the most: Eugenio Carpizo-Ituarte, Curt Fiedler, Lucia Gutierrez, Tim Male, v Javier Mendez, Dwayne Minton, Teresa Restom, Winnie Roberts, Andre Seale, and Reuben Wolff. I thank Tom Duda, Claudia Farfan, Keith Fong, and Tricia Murata for help in the molecular lab. I also thank Chris Meyer (FLMNH) for help in designing cowrie-specific primers and sharing protocol tips. Many friends in Brazil and elsewhere provided encouragement and friendship via email, and almost made me forget I lived in one ofthe most isolated places in the world. Finally, I thank my family in Brazil and my in-laws in Hawaii for their support and encouragement, and my wife Heather for her love, patience and for our daughter Olivia and another baby girl due soon. Below are acknowledgments pertinent to each chapter: Chapter 2 I thank Chris Meyer (FLMNH) for planting the seed for some ofthe ideas presented here, Rebecca Price (Univ. Chicago) for discussion, and John Earle and Wes Thorsson (Hawaii), and Scott Johnson (Kwajalein) for photographs oflive cowries. Chapter 3 I thank E. Alison Kay and Alan J. Kohn (University ofWashington) for discussion and suggestions, and E. A. Kay and Hugh Bradner (Scripps Institution) for donation ofmaterial. I also thank the many donors ofCypraeidae acknowledged by Bradner and Kay (1996) in their Atlas ofCowrie Radulae. Many ofthe odontophores illustrated here, including several from rare species, were obtained from the specimens vi used previously in Bradner and Kay (1996). I thank John Earle for the donation ofa rare specimen ofLyncina leucodon. Most ofthe odontophores in the Cribrarula cribraria complex were obtained from dried specimens from the museums listed above. I thank Paul Callomon (ANSP) for helping with the donation ofodontophore stubs to the museum. I am grateful to H. Gert de Couet, Department ofZoology, University ofHawai'i for allowing me to perform the experimental part ofthis work in his molecular laboratory. The assistance ofTina M. Weatherby with SEM at the University ofHawai'i Biological Electron Microscope Facility (BEMF) and a grant to BEMF (#RR1AI03061) which partially subsidized SEM costs are much appreciated. I also thank James Parrish and Lucy Kida, Cooperative Research Unit, University ofHawai'i for access to a digital camera. This project could not have been possible without the financial support from the Hawaiian Malacological Society, for which I say Mahalo Nui Loa! Chapter 4 I thank E. Alison Kay and Carole Hickman for suggestions on the manuscript. Some material was donated by Kay and Hugh Bradner (Scripps Institution), some loaned from several ofthe museums listed above. I also thank Tina M. Weatherby for help with SEM at BEMF; and James Parrish and Lucy Kida made possible the use ofa digital camera and dissecting scope used to produce some ofthe figures shown here. VB ChapterS I thank Ian Loch, AMS, for processing the specimen loan, responding to inquiries and his efforts to locate additional specimens ofC. gravida Moretzsolm 2002. I am thankful to Felix Lorenz, Institut fur Allgemeine und Spezielle Zoologie, Germany, for access to literature and additional information on the recently described C. garciai, and Chris Meyer, Florida Museum ofNatural History, for designing cowrie-specific primers and useful tips and molecular biology protocols. Chapter 6 This review could not have been done without access to museum collections. I thank the curators and collection managers from the museums listed above for assistance with arranging visits and specimen loans, and giving permission to illustrate some ofthe specimens. I thank Jackie van Goethem, Royal Belgian Institute ofNatural Sciences, Brussels, Belgium; Bernard Metivier and Philippe Bouchet, Museum National d'Histoire Naturelle, Paris, France for helping with locating specimens, and Kathie Way and Amelia Campbell, Natural History Museum, London, UK, for information and photographs of types in the British Museum. I am indebted to the many shell collectors and biologists listed above who contributed specimens, literature and information. I thank Wallace Holznagel (Smithsonian Institution), Bob Kinzie (University of Hawaii), Alan Kohn (University ofWashington, Seattle), Jose Leal (Bailey-Matthews Museum, Florida), Felix Lorenz (Institut fur Allgemeine und Spezielle Zoologie, Germany), Chris Meyer (FLMNH), and Charlie Sturm Jr. (CMNH) for helpful viii information, suggestions and discussions. I also wish to acknowledge with thanks Sue Monden for producing line drawings of shells, and Wes Thorsson and Scott Johnson who generously made their excellent photographs oflive specimens available. I thank Lindsey Groves, Chris Meyer, and two anonymous reviewers, for their detailed comments, which considerably improved the manuscript. IX ABSTRACT The goal ofthis dissertation is to explore novel and non-traditional taxonomic characters that may be useful for mollusks, and combine them with radular and shell characters to carry out a taxonomic review ofthe genus Cribrarula (Cypraeidae). Shells in this genus have conspicuous dorsal spots, but like odler cowries, lack sculpture, spines, and other shell characters commonly used in gastropod taxonomy. For these reasons, cowrie shells are considered uninformative. The study ofdorsal spots (DS) and related characters suggest that at least in this complex, the dorsal spots may represent a record ofthe mantle papillae. If the hypothesis is correct, then DS may provide information on the soft parts that previously was only available from the study oflive or preserved specimens. Each species in the complex has a species-specific range ofDS, marginal spots, and allied characters, thus suggesting that they may be useful in distinguishing species in the complex. The odontophore cartilage provides support for the radula and attachment for the muscles responsible for feeding. Although intimately connected to the radula and known since the 1800's, the taxonomic value ofthe odontophore