Durham E-Theses The palaeobiology and systematics of some Jurassic bryozoa Taylor, P. D. How to cite: Taylor, P. D. (1977) The palaeobiology and systematics of some Jurassic bryozoa, Durham theses, Durham University. Available at Durham E-Theses Online: http://etheses.dur.ac.uk/8271/ Use policy The full-text may be used and/or reproduced, and given to third parties in any format or medium, without prior permission or charge, for personal research or study, educational, or not-for-prot purposes provided that: • a full bibliographic reference is made to the original source • a link is made to the metadata record in Durham E-Theses • the full-text is not changed in any way The full-text must not be sold in any format or medium without the formal permission of the copyright holders. Please consult the full Durham E-Theses policy for further details. Academic Support Oce, Durham University, University Oce, Old Elvet, Durham DH1 3HP e-mail: [email protected] Tel: +44 0191 334 6107 http://etheses.dur.ac.uk THE PALEOBIOLOGY AND SYSTEMATICS OF SOME JURASSIC BRYOZOA by P.D. Taylor Van Mildert College A thesis presented for the degree of Doctor of Philosophy in the University of Durham Volume 1 - Text Department of Geological Sciences, University of Durham. September, 1977 The copyright of this thesis rests with the author. No quotation from it should be published without his prior written consent and information derived .' V.!i7 ' ''?i from it should be acknowledged. ii ABSTRACT Morphological studies of some non-fasciculate Jurassic tubuloporinids and comparative studies of other cyclostomatous Bryozoa have enabled aspects of their palaeobiology to be elucidated and their systematics to be revised. Skeletal ultrastructure has been surveyed in a number of species and its relevance as a taxonomic character assessed. Zooid structure and variation are considered. Factors influencing ontogenetic and astogenetic zonation of colonies are suggested and the functional significance of zonation is postulated. The functional morphology and genesis of tubuloporinid heterozooids have been elucidated. Quantitative studies have shown that ecophenotypic within-colony zooidal variation is high and tends to dominate between-colony comparisons of zooecium size. Differences in colony growth-form are a product of differing styles of zooecial budding, forms of budding zones, and positions of budding loci. Unilamellar, bilamellar, multilamellar, non-lamellar, and composite (Terebellaria) patterns of colony growth are described. 8 Jurassic tubuloporinid growth-forms are recognised and their ecological significance assessed by functional morphological inference and comparison with living analogues. Strength and stability, integrated zooid feeding, and iii differential modes of resource utilization are shown to be reflected by colony growth-form. The ecological succession of bryozoan growth-forms is inferred. Three families of non-fasciculate tubuloporinids are recognised in the Jurassic; Stomatoporidae, Multisparsidae and Plagioeciidae. Emended diagnoses are given of 9 genera together with a key to the identification of all accepted genera and systematic descriptions of 15 species including two new species, Reptomultisparsa tumida and Reptoclausa porcata. The palaeoecology of Jurassic bryozoans has been examined and their distribution is shown to be largely facies controlled whilst species exhibit extended longevities. ACKNOWLEDGEMENTS I wish to thank my supervisors Dr. G.P. Larwood and Miss P.L. Cook (BMNH) for their encouragement, advice and constructive criticism throughout the course of this project. I have gained much from discussions with Drs. K. Brood (Naturhistoriska Riksmuseet, Stockholm), P.J. Hayward (University College of Swansea),H.C. Jenkyns (University of Oxford), J.R. Nudds (Trinity College, Dublin), T.J. Palmer (University of Oxford), B.R. Rosen (BMNH) and J.R. Senior (University of Durham), and from correspondence with Dr. G. lilies (University of Karlsruhe) and Dr. R.L. Anstey (Michigan State University). I also wish to thank R.W. Furness (Department of Zoology, University of Durham) with whom the study of zooidal size variation in Stomatopora colonies was undertaken and J. Gould who collected the specimens for • this study. Loans of specimens were very kindly arranged through Miss J. Darrell, Mr. R. Wise and Mr. P.J. Chimonides (BMNH), Dr. S. Tillier and Dr. E. Buge (MNHN), H.P. Powell (OUM), Miss B. Pyrah (YM) and Mr. G. Spalton (RUGD). I am extremely grateful to Mrs. H. Winn and Mrs. L. Mines for typing this thesis, to Mr. G. Dresser and Mr. J. Clayton for photographic work, and to other members of the technical staff for their valued assistance. V For facilities in the Department of Geological Sciences I wish to extend my gratitude to Professors M.H.P. Bott and G.M. Brown, and for the provision of a research studentship I am indebted to the Natural Environment Research Council of Great Britain. Finally, I thank my wife Patricia for the patience and tolerance she displayed during fieldwork and the preparation of this thesis, and for her very welcome help with proof reading. CONTENTS VOLUME 1 Page ABSTRACT ii ACKNOWLEDGEMENTS iv CONTENTS vi LIST OF TABLES ix CHAPTER 1 INTRODUCTION 1 CHAPTER 2 MATERIALS AND METHODS 7 CHAPTER 3 CYCLOSTOME ANATOMY 21 CHAPTER 4 SKELETAL ULTRASTRUCTURE 41 CHAPTER 5 ZOOIDAL ONTOGENY 51 CHAPTER 6 ASTOGENETIC ZOOIDAL VARIATION 73 CHAPTER 7 ZOOIDAL POLYMORPHISM 87 CHAPTER 8 WITHIN COLONY ENVIRONMENTAL ZOOIDAL VARIATION 118 CHAPTER 9 BETWEEN COLONY AUTOZOOIDAL VARIATION 133 CHAPTER 10 COLONY GROWTH-FORM 145 CHAPTER 11 COLONY GROWTH 161 CHAPTER 12 UNILAMELLAR PATTERNS OF COLONY GROWTH 177 CHAPTER 13 BILAMELLAR PATTERNS OF COLONY GROWTH 195 CHAPTER 14 MULTILAMELLAR PATTERNS OF COLONY GROWTH 199 CHAPTER 15 NON-LAMELLAR PATTERNS OF COLONY GROWTH 213 vii Page CHAPTER 16 COMPOSITE PATTERN OF COLONY GROWTH ; TEREBELLARIA 221 CHAPTER 17 FUNCTIONAL MORPHOLOGY AND MECHANICAL PROPERTIES OF ZOARIA 242 CHAPTER 18 EXTRAZOOIDAL FEEDING CURRENTS AND ZOARIAL MORPHOLOGY 259 CHAPTER 19 THE FUNCTIONAL MORPHOLOGY OF RESOURCE UTILIZATION 277 CHAPTER 20 COLONIALITY IN THE CYCLOSTOMATA 2 93 CHAPTER 21 PALAEOECOLOGY 308 CHAPTER 22 SYSTEMATICS 339 Suborder Tubuloporina 353 A key to the non-fasciculate genera of Jurassic tubuloporinids 356 Family Stomatoporidae 359 Family Multisparsidae 363 Genus Reptomultisparsa 366 Reptomultisparsa incrustans 369 Reptomultisparsa tumida 376 Genus Reptoclausa 379 Reptoclausa porcata 381 Genus Collapora 388 Collapora straminea 391 Collapora microstoma 400 Collapora tetragona 411 Family Plagioeciidae 417 Genus Hyporosopora 422 Hyporosopora typica 424 Hyporosopora parvipora 430 viii. Page Hyporosopora portlandica 436 Hyporosopora sauvagei 440 Genus Mesenteripora 445 Mesenteripora undulata 448 Genus Reticulipora 457 Reticulipora dianthus 459 Genus Entalophora 466 Entalophora annulosa 468 Genus Mecynoecia 474 'Mecynoecia' bajocina 477 Genus Terebellaria 482 Terebellaria ramosissima 483 CHAPTER 2 3 CONCLUSION 491 REFERENCES 494 VOLUME 2 FIGURES PLATES APPENDIX 1 FIELD LOCALITIES xi APPENDIX 2 PUBLICATIONS lix ix LIST OF TABLES Following page Table 1. Comparative features of single-walled and double-walled cyclostomes. 35 Table 2. Comparison of autozooecial dimensions and kenozooecial concentration between different zones of astogenetic repetition in a Collapora microstoma colony. 78 Table 3. Zooecial frontal wall length in zoaria of Stomatopora bajocensis and S.dichotomoides. 81 Table 4. Zooecial frontal wall width in zoaria of Stomatopora bajocensis and S.dichotomoides. 81 Table 5. Correlations between zooecial frontal wall length and zooecial generation in zoaria of Stomatopora bajocensis and S.dichotomoides. 83 Table 6. Correlations between zooecial frontal wall width and zooecial generation in zoaria of Stomatopora bajocensis and S.dichotomoides. 83 Table 7. Comparison between the zooecial generation number marking the onset of astogenetic repetition determined by the linear regression method and that determined subjectively. 84 Table 8. Between colony variation in gono- zooecial characters in Mesenteripora undulata from Luc-sur-mer. 110 Table 9. Environmental variation in zooecial dimensions in Stomatopora colonies. 125 Table 10. Data used in the F tests comparing the contributions of within and between colony variance to total variance in Stomatopora. 127 X Following page Table 11. Analysis of variation within and between generations in Stomatopora colonies. 127 Table 12. F tests comparing the contributions of within and between colony variance to total variance in Reptomultisparsa incrustans. 135 Table 13. F tests comparing the contributions of within and between colony variance to total frontal wall length variance in Mesenteripora undulata. 136 Table 14. F tests comparing the contributions of within and between colony variance to total variance in Terebellaria ramosissima. 137 Table 15. Between colony CVs for autozooecial characters in the tubuloporinids studied. 138 Table 16. Zoarial morphological features indicative of the presence of an extrazooidal water current system 263 Table 17. Upper Bajocian and Bathonian stratigraphy of Calvados, Normandy. XI Table 18. Bathonian stratigraphy of southern England. XXXI1 Table 19. Inferior Oolite stratigraphy of the Cotswolds. xxx ix CHAPTER 1 INTRODUCTION The Bryozoa are a phylum of aquatic filter-feeding organisms which are exclusively colonial and usually sessile. Most authors recognise 3 bryozoan classes; Phylactolaemata, Gymnolaemata and Stenolaemata.