The Evolution of the Cephalaspidea (Mollusca: Gastropoda) and Its Implications to the Origins and Phylogeny of the Opisthobranchia Terrence Milton Gosliner
Total Page：16
File Type：pdf, Size：1020Kb
University of New Hampshire University of New Hampshire Scholars' Repository Doctoral Dissertations Student Scholarship Spring 1978 THE EVOLUTION OF THE CEPHALASPIDEA (MOLLUSCA: GASTROPODA) AND ITS IMPLICATIONS TO THE ORIGINS AND PHYLOGENY OF THE OPISTHOBRANCHIA TERRENCE MILTON GOSLINER Follow this and additional works at: https://scholars.unh.edu/dissertation Recommended Citation GOSLINER, TERRENCE MILTON, "THE EVOLUTION OF THE CEPHALASPIDEA (MOLLUSCA: GASTROPODA) AND ITS IMPLICATIONS TO THE ORIGINS AND PHYLOGENY OF THE OPISTHOBRANCHIA" (1978). Doctoral Dissertations. 1197. https://scholars.unh.edu/dissertation/1197 This Dissertation is brought to you for free and open access by the Student Scholarship at University of New Hampshire Scholars' Repository. It has been accepted for inclusion in Doctoral Dissertations by an authorized administrator of University of New Hampshire Scholars' Repository. For more information, please contact [email protected]. INFORMATION TO USERS This material was produced from a microfilm copy of the original document. While the most advanced technological means to photograph and reproduce this document have been used, the quality is heavily dependent upon the quality of the original submitted. The following explanation of techniques is provided to help you understand markings or patterns which may appear on this reproduction. 1.The sign or "target" for pages apparently lacking from the document photographed is "Missing Page(s)". If it was possible to obtain the missing page(s) or section, they are spliced into the film along with adjacent pages. This may have necessitated cutting thru an image and duplicating adjacent pages to insure you complete continuity. 2. When an image on the film is obliterated with a large round black mark, it is an indication that the photographer suspected that the copy may have moved during exposure and thus cause a blurred image. You will find a good image of the page in the adjacent frame. 3. When a map, drawing or chart, etc., was part of the material being photographed the photographer followed a definite method in "sectioning" the material. It is customary to begin photoing at the upper left hand corner of a large sheet and to continue photoing from left to right in equal sections with a small overlap. If necessary, sectioning is continued again — beginning below the first row and continuing on until complete. 4. The majority of users indicate that the textual content is of greatest value, however, a somewhat higher quality reproduction could be made from "photographs" if essential to the understanding of the dissertation. Silver prints of "photographs" may be ordered at additional charge by writing the Order Department, giving the catalog number, title, author and specific pages you wish reproduced. 5. PLEASE NOTE: Some pages may have indistinct print. Filmed as received. University Microfilms International 300 North Zeeb Road Ann Arbor, Michigan 48106 USA St. John's Road. Tyler's Green High Wycombe, Bucks, England HP10 8HR 7624553 GOSLINER# TERRENCE MILTON THE EVOLUTION OF THE CEPHALASPlDEA IIMOLLUSCAiGASTROPOOA) AND ITS IMPLICATIONS TO THE ORIGINS AND PHYLOGENY OF THE opisthobranchia , UNIVERSITY OF NEW HAMPSHIRE* PH.D., 1978 University Microfilms International 300 n . z e e b r o a d , a n n a r b o r ,m i 4 8 io 6 © 1978 TERRENCE MILTON GOSLINER ALL RIGHTS RESERVED THE EVOLUTION OF THE CEPHALASPIDEA (MOLLUSCA: GASTROPODA) AND ITS IMPLICATIONS TO THE ORIGINS AND PHYLOGENY OF THE OPISTHOBRANCHIA by TERRENCE M. GOSLINER A.B., University of California, 1972 M.S., University of Hawaii, 1973 A THESIS Submitted to the University of New Hampshire In Partial Fulfillment of The Requirements for the Degree of Doctor of Philosophy Graduate School Department of Zoology May, 1978 This thesis has been examined and approved. tfleiMu A.. Thesis director, Larry Harris, Assoc. Prof. of ZoologJ ^ A r t h u r C. Borror, Prof. of Zoology M. Palfricia Morse, Assoc. Prof. of Biology, Northeastern University </[s> Robert Robertson, Pilsbry Chair of Malacology Academy of Natural Sciences, Philadelphia Emery F. Sv/an/ Prof. of Zoology e ACKNOWLEDGEMENTS I am grateful for the support and assistance of my doctoral committee, Drs. Arthur C. Borror, M. Patricia Morse, Robert Robertson, and Emery F. Swan. I especially thank my thesis advisor, Dr. Larry G. Harris, for his guidance, challenging ideas and friendship. Discussion with Drs. Ruth Turner and Michael T. Ghiselin provided valuable input to my research efforts. I would like to thank the following individuals for their assistance in obtaining and collecting preserved and living specimens: Dr. Hans Bertsch, Chaminade College, Honolulu, Hawaii; Dr. Philippe Bouchet, Museum National de Histoire Naturelle, Paris, France; Mr. Dustin Chivers, California Academy of Sciences, San Francisco; Dr. George Davis, Academy of Natural Sciences, Philadelphia, Pennsylvania; Ms. Bonnie J. Gosliner; Mr. Michael Gosliner, Sleepy Hollow Institute of Technology, San Anselmo, California; Ms. Virginia Gosliner; Dr. Willard Hartman, Peabody Museum, Yale University, New Haven, Connecticut; Dr. June Harrigan, Marine Biological Laboratory, Woods Hole, Massachusetts; Dr. Larry G. Harris, University of New Hampshire; Ms. Pele Harris; Mr. Alan Hulbert, University of New Hampshire; Dr. Alan Kuzirian, University of New Hampshire; Dr. Eveline Marcus, University of Sao Paulo, Brazil; Ms. Kaniaulono Meyer, University of Cincinnati, Ohio; Ms. Sandra Millen, University of British Columbia, Vancouver; Mr. David Mulliner; Dr. M. Patricia Morse, Northeastern University, Boston, Massachusetts; Mr. Frank Perron, University of Hawaii, Honolulu; Dr. Michael Rex, University of Massachusetts, Boston; Mr. Brian Rivest, University of Washington, Seattle Dr. Robert Robertson, Academy of Natural Sciences, Philadelphia, Pennsylvania; Mr. Gale Sphon, Los Angeles County Museum, California; iii Dr. Ruth Turner, Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts; and Mr. Gary C. Williams, Oak Manor Agricultural Station, Fairfax, California. My deepest appreciation goes to my wife, Bonnie, who assisted in all stages of my work, particularly in the preparation of the dissertation. iv TABLE OF CONTENTS List of Tables................................................ vii List of Plates................................. viii Abstract...................................................... xiii Common Introduction........................................... 1 Chapter I: A Morphological Comparison of Utriculastra and Cylichnella (Opisthobranchia: Scaphandridae).................. 3 Abstract...................................................... 3 Introduction.................................................. 3 Materials and Methods......................................... 4 Results............................ 5 Discussion.................................................... 10 Literature Cited.............................................. 15 Chapter II: Systematics and Phylogeny of the Aglajidae (Opisthobranchia: Mollusca)................................... 23 Introduction.................................................. 23 Methods. ................................................... 25 Aglaja Reiner, 1807........................................... 25 Discussion of Aglaj a ..................... 26 Discussion of Species Placed in Aglaj a ........................ 30 Chelidonura A. Adams, 1850.................................... 34 Discussion of Chelidonura Adams, 1850........................ 35 Discussion of Chelidonura species............................ 37 Melanochlamys Cheeseman, 1881................................. 40 Discussion of Melanochlamys Cheeseman, 1881.................. 41 Discussion of Species Placed in Melanochlamys................. 42 Navanax Pilsbry, 1895......................................... 44 Discussion of Navanax Pilsbry, 1895.......................... 45 Discussion of Navanax species................................. 46 Phil inop sis Pease, 1860....................................... 52 Discussion of Philinopsis Pease, 1860........................ 54 Discussion of Species Placed in Philinopsis................... 55 Discussion of Pseudophiline Habe, 1976 and Philinorbis Habe, 1950........................................................ 57 Discussion of Odontaglaja Rudman, 1978........................ 58 Evaluation of Morphological Characters........................ 58 Systematics and Phylogeny..................................... 69 Acknowledgements.............................................. 72 References.................................................... 72 Chapter III: Origins and Relationships of Primitive Members of •the Opisthobranchia (Mollusca: Gastropods).................... 91 v 0 ;■! ; Introduction.................................................. 91 Material Examined............................................. 92 Results and Discussion........................................ 92 I. Definition of Opisthobranchia; included and excluded taxa.... 92 II. Common Ancestry with the Pulmonata........................ 96 Ilia. Origin of the Opisthobranchia and Pulmonata from Archaeogastropods........................................... 98 Illb. Origins of the Opisthobranch-pulmonate clade from Rissoacean-Cerithiaceans.................................... 101 IIIc. Origins of the Opisthobranch-pumonate clade from Littorinacea..................................... 106 ; IVa. Relative Primitiveness of Acteon and other Acteonidae 109 ; IVb. The Primitiveness