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Photographs included in ttie original manuscript have been reproduced xerographically in this copy. Higher quality 6” x 9” black arxt white photographic prints are available for any photographs or illustrations appearing in this copy for an additional charge. Contact UMI directly to order. Bell & Howell Information and Learning 300 North Zeeb Road, Ann Arbor, Ml 48106-1346 USA 800-521-0600 UMI* Phylogeny of Vestimentiferan Tube Worms by Anja Schulze Diplom, University of Bielefeld, 1995 A Dissertation Submitted in Partial Fulfillment of the Requirements for the Degree of DOCTOR OF PHILOSOPHY in the Department of Biology We accept this dissertation as conforming to the required standard Dr. '/TTunniphffe, Supervisor (Department of Biology) Dr. IvTR. Page (Ekpartment of Biology) Dr. R. IX Burke (Department of Biology and Department of Biochemistrv'/Microbioloav) _____________________________________________________________ Dr. C (School of Earth and Ocean Science) Dr. D. McHugh, External Examiner (ColgateUniversity) © Anja Schulze, 2000 University of Victoria Ail rights reserved. This dissertation may not be reproduced in whole or in part, by photocopying or other means, without the permission of the author. Abstract Supervisor; Dr. Verena Tunnicliffe Vestimentifera inhabit hydrothermal vents, cold-water seeps and other marine reducing habitats. The objectives of this study were to analyse phylogenetic relationships among the extant species and their affinities to perviate and moniliferan Pogonophora and Polychaeta. The phylogeny was reconstructed using morphological characters to test phylogenetic hypotheses based on molecular data. Morphological characters were partly extracted from the literature and partly gained throughout study of gross morphological and anatomical investigations and light, transmission and scanning electron microscopy. Three aspects of morphology were examined in detail in nine vestimentiferan species. The excretory system differs among the vestimentiferan species in the number of excretory pores, absence/presence of excretory papillae and grooves and shape of the excretory ducts. The anatomy of the excretory system resembles that shared by the polychaete families Serpulidae. Sabellidae and Sabellariidae. Chaetal ultrastructure and chaetogenesis show patterns similar to uncini in polychaetes. Contraiy to published accounts, the septa dividing the opisthosomal segments only bear musculature on their posterior faces. A rudimentary gut and anus are present in opisthosomes of specimens up to adult size. The blood vascular system includes an intravasal body in the dorsal vessel with ultrastructural characteristics similar to intravasal tissue in Terebellidae, Ampharetidae, Flabelligeridae and Serpulidae, and is probably involved in hemoglobin production. Hemocytes were detected in many blood vessels, most of them attached to the vascular lamina. The sinus valvatus is a specialised region of the anterior ventral vessel, apparently unique to vestimentiferans. The wall of the dorsal vessel is formed by myoepithelial cells, representing a coelomyarian type of double obliquely striated musculature. Phylogenetic analyses including a total of 17 vestimentiferan species and three perviate species as outgroups support molecular interpretations that the vestimentiferan species inhabiting basalt-hosted vents of the Eastern Pacific represent a derived monophyletic clade. According to the reconstructed phylogeny, the ancestral habitat of Vestimentifera was deep-water sedimented vent sites in the Western Pacific. Analysis of the relationships among Pogonophora and six polychaete families placed Ill Pogonophora at the base of a clade including Sabellidae, Serpulidae and Sabellariidae. The Oweniidae represent the sister group to this clade. Examiners: Dr. V. Tunnicliffe, supervisor (Department of Biology) :_ Dr. L. 1^ Page (D eg^m ent of Biology) Dr. R. D. Burke (Department of Biology and Department of Biochemistry/Microbiology) Dr. C. R. chool of Earth and Ocean Science) Dr. D. McHugh, External Examiner (Colgate University) IV Table of Contents page Title Page i Abstract ii Table of Contents iv List of Tables vi List of Figures viii Acknowledgments x Chapter 1: Introduction 1 Literature Cited 8 Chapter 2: Comparative Anatomy of Excretory Organs in 12 Vestimentiferan Tube Worms (Pogonophora, Obturata) Abstract 12 Introduction 12 Material and Methods 13 Results 15 Discussion 34 Literature Cited 37 Chapter 3: The Opisthosome: Ultrastructure of Opisthosomal Chaetae 39 in Vestimentifera (Pogonophora, Obturata) and Implications for Phylogeny Abstract 39 Introduction 40 Material and Methods 43 Results 45 Discussion 56 Literature Cited 59 Appendix: Additional Characters of the Opisthosome 63 Chapter 4: The Blood Vascular System: Histological and 71 Ultrastructural Characterisation with an Emphasis on the Intravasal Body Introduction 71 Material and Methods 74 Results 74 Discussion 89 Literature Cited 93 Chapter 5: Phylogenetic Relationships among Vestimentiferans from 97 Hydrothermal Vents and Cold-Water Seeps Introduction 97 Methods 102 Results 120 Discussion 133 Literature Cited 143 Appendix 5: Characters Used in Phylogenetic Analysis 150 Chapter 6: Affinities bet^veen Pogonophora and Polychaeta 157 Introduction 157 Methods 159 Results 166 Discussion 168 Literature Cited 176 Appendix 6.1 : Characters for Reconstruction of the Ancestral 184 Pogonophoran Appendix 6.2: Characters for Family Level Analysis 188 Chapter 7: Summary 192 Literature cited 192 Appendix: Vestimentiferans (Pogonophora) in the Pacific and Indian 197 Oceans: a new genus from Lihir Island (Papua New Guinea) and the Java Trench, with the first report of Arcovestia ivanovi from the North Fiji Basin List of Tables page Table 1 : History of classification of Vestimentifera and Pogonophora 4 Table 2: Differences among eight vestimentiferan species with regard to their 19 excretory systems Table 3 .1 : Presence of uncini and rod-shaped chaetae in juvenile and adult 42 Pogonophora Table 3.2: Material examined, fixation and use 44 Table 3.3: Proportion of total number of segments and chaetal rows per 48 segment in ten species of Vestimentifera Table 3.4: Comparison of uncini in three polychaete orders and Pogonophora 57 Table 4.1 : Geographic origin, body regions examined, diameter of specimen in 75 vestimentaJ region and maximum diameter of intravasal body Table 4.2: Results of different staining procedures in Riftia pachyptila 79 Table 5.1 : Vestimentiferan species described or mentioned in the literature to 98 date, their inclusion in the present study, general geographic area, specific sites, depth ranges and type of habitat Table 5.2: Characters and character states used in the conventional coding and 104 multistate coding analysis Table 5.3: Species level dataset, multistate coding 108 Table 5.4: Species level dataset, conventional coding 109 Table 5.5: Genus level dataset, multistate coding 110 Table 5.6: Genus level dataset, conventional coding 111 Table 5.7: Coding strategies illustrated with an example from the present 112 analysis Table 5.8: Species ranked according to mean obturaculum/vestimentum length 115 Table 5.9: Species ranked according to mean vestimentum length/diameter 115 Table 5.10: Species ranked according to their number of branchial lamellae 118 vu Table 5 .11 : Species ranked according to mean number of opisthosomal 118 segments Table 5.12: Summary of results of the species level analyses 121 Table 5.13: Summary of results of the species level analyses excluding Alaysia 121 spiralis and Lamellibrachia victori Table 5.14: Summary of results of the genus level analyses 122 Table 5.15: Summary of results of the genus level analyses excluding Alaysia 122 Table 6.1 : Species used in initial analysis and their traditional classification 160 Table 6.2: Characters and character states used in initial analysis 162 Table 6.3: Dataset for reconstruction of the ancestral pogonophoran 163 Table 6.4: Characters and character states used in family level analysis 164 Table 6.5: Dataset for family level analysis, excluding larval characters 165 Table 6.5: Larval characters 165 Table A1 : Dimensions of Paraescarpia echinospica sp. nov. gen. nov (type 207 material) Table A2: Characteristics of vestimentiferan genera 217 VIII List of Figures page Figure 1 2 Figure 2.1 17 Figure 2.2 22 Figure 2.3 25 Figure 2.4 27 Figure 2.5 29 Figure 2.6 33 Figure 3.1 47 Figure 3.2 50 Figure 3.3 52 Figure 3.4 55 Figure 3.5 65 Figure 3.6 68 Figure 3.7 70 Figure 4.1 77 Figure 4.2 81 Figure 4.3 83 Figure 4.4 86 Figure 4.5 88