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Grania (Annelida, Clitellata, Enchytraeidae) of the Bermuda Islands

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Grania (Annelida, Clitellata, Enchytraeidae) of the Bermuda Islands SYSTEMATICS AND BIOLOGY OF GRANlA (ANNELI DA: CLITELLATA: ENCHYTRAEIDAE) OF THE BERMUDA ISLANDS Jan Maureen Locke A thesis submitted in conforrnity with the requirements for the degree of Master of Science Graduate Department of Zoology University of Toronto O Copyright by Jan Maureen Locke 1998 National tibraw Bibliothwue nationale du Canada Acquisitions and Acquisitions et Bibliogiaphic Sentices services bibliographiques 395 Wellington Street 395, nie Wellington OnawaON K1AW OnawaON K1AON4 Canada Canada The author has granted a non- L'auteur a accordé une licence non exclusive Licence dowing the exclusive permettant à la National Library of Canada to Bibliothèque nationale du Canada de reproduce, ioan, distribute or seIl 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/film, 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 fiom it Ni la thèse ni des extraits substantiels may be printed or othewise de celle-ci ne doivent être imprimés reproduced without the author's ou autrement reproduits sans son permission. autorisation. Systernatics and Biology of Grania (Annelida, Clitellata, Enchytraeidae) of the Bermuda Islands MSc 1998 Jan M. Locke Department of Zoology, University of Toronto The diversity and distribution of the marine enchytraeid genus Grania present within intertidal and subtidal coastal habitats of the Atlantic island of 8ermuda are investigated. Two new species, Grania laxarta and Grania hylae, are recorded and Grania bermudensis and Grania amerkana are redescribed from Benuda, Florida and the Caribbean. These recent descriptions incorporate the new taxonomic character information of bipartite sac male apparatus and the anterior cerebral statocyst. This statocyst is the first description within clitellate annelids. These new informative characteristics were combined with a compilation of familiar characters for the genus to produce a dichotomous key for eight species of Gfania and two species of the genus Randidfilus from the western Atlantic and Caribbean. Grania species from these areas were also used in a phylogenetic study to determine possible evolutionary relationships among the Bermudian species. An evolutionary hypothesis of dispersal by currents, the Gulf Stream in particular, was suggested for the species of Grania that occupy Bermuda, due to its volcanic, oceanic island history. The combination of good taxonomic information with characteristically small ranged species enable Grania to be a rnodel taxon for zoogeographic studies of benthic marine invertebrates. ACKNOWLEDGMENTS The encouragement and patience given to me by my advisor and friend Kathryn Coates, is deeply appreciated. I was extremely fortunate to be surrounded by her wealth of expert knowledge. I do hope some of it nibbed off. Dr. Doug Currie, my adjunct supervisor provided valuable assistance and was a great contact in Toronto during my period of research abroad. I am also grateful to MI. Don Stacey of the Royal Ontario Museum, for his patience and pointing me in the right direction when it came to "worm work. My research was conducted at the Bermuda Biological Station for Research, Inc. as part of the BBSR graduate intern program. Thank you to the faculty and staff of BBSR for their kindness during my time there. Several individuals contributed continued support during this work: my parents Donna and Wallace Locke, my friends Michael Stevens, Jason Ritter, Jarney Penney-Ritter, Ruth Kelty and Astrid Schnetzer for her German translations and assistance in the final preparations. I am also grateful to Mr. Rudy Villadiego for electron microscopy technical assistance and Hyla Tibbitts and Linda Nelson who provided detailed illustrations of Grania that I truly appreciated. I gratefully acknowledge the financial support provided by, University of Toronto Open Masters Fellowship 1996-1998; The Canadian Associates of the Bermuda Biological Station; The Roger Thayer Stone Fellowship, The Munson Foundation and Hank Trapido-Rosenthal through the Bermuda Biological Station for Research. Support and funding for research facilities were also provided by National Science Foundation grants # DEB-9615211 and # DBI 9724335. iii For Donna and Wallace TABLE OF CONTENTS TlTLE PAGE .................................................................................................................... i ABSTRACT ......................~............................................................................................. ii ... ACKNOWLEDGEMENTS ................................................................................................ III DEDICATION ................................................................................................................. iv TABLE OF CONTENTS .................... ,., ....., ...................................................................... v ... LIST OF TABLES .~......................................................................................................... viii LIST OF FIGURES ....................................................................................................... ix 1 . GENERAL INTRODUCTION ..............................................................................1 The Genus ....................................................................................................5 History ..........................................................................................................6 Morphology ..................................... ...........................................................13 Habitat and Distribution ............................................................................17 Ecology ....~...~..........................................................................................18 Reproductive Biology ............................................................................. 18 Copulation ................................................................................................20 Taxonomie Difficulties ...~.............................................................................21 Objectives ...................................................................................................22 References .......................... ......... .......................................................... 24 2. REDESCRIPTIONS OF GRANlA AMERCANA. G. BERMUDENSIS AND DESCRIPTIONS OF TWO NEW SPECIES OF GRANIA (ANNELIDA: CLITELîATA: ENCHYTRAEIDAE) FROM BERMUDA .............................. 31 Introduction ............................. ... ............................................................33 Materials and Methods .............................................................................. 36 Results ...................................................................................................... 41 New Species Descriptions ....................................................................41 Granja laxarla ...................................................................................41 Grania hylae ....................................................................................51 Species Redescriptions ........................................................................61 Grania bermudensis ........................................................................61 Grania americana ..........................................................................71 Discussion .................................... ..,. .......................................83 References .............................................................................................88 3. ULTRASTRUCTURAL STUDIES OF THE STATOCYST OF THE MARINE ENCHYTRAEID GRANlA AMERCANA (ANNELIDA: CLITELLATA) ............. 92 Abstract ....................................................................................................92 Introduction .................................................................................................92 Materials and Methods ................................................................................94 Results .......................................................................................................95 Light Microscopy ................................................................................ 95 Transmission Electron Microscopy ................................................... 98 Discussion ............................................................................................... 109 References .............................................................................................118 4 . A KEY TO THE SPECIES OF GRANlA (ENCHYTRAEIDAE: CLITELLATA) OF EASTERN NORTH AMERICAN AND NEARBY COASTAL AND CARIBBEAN POPULATIONS ............................................................................................ 125 Abstract ................................................ ................................................. 125 Introduction ............................................ ........................................... 125 Grania of the Northwest Atlantic ............................................. 126 Recognition of Grania .......................... ... ...................................... 131 Definitions and Diagnostic Characters ...............................................132 Materials
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