Kelso-Maguire Thesis 2020.Pdf
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An investigation into the Bryozoa of Ireland including the distributional patterns of the marine Bryozoa of Ireland over the last 150 years By Antoinette Kelso-Maguire A thesis submitted to Trinity College Dublin in fulfilment of the requirements for the degree of Doctor of Philosophy 2020 I declare that this thesis has not been submitted as an exercise for a degree at this or any other university and it is entirely my own work. I agree to deposit this thesis in the University’s open access institutional repository or allow the library to do so on my behalf, subject to Irish Copyright Legislation and Trinity College Library conditions of use and acknowledgement. TABLE OF CONTENTS Acknowledgements .......................................................................................................... iv List of figures .................................................................................................................... v List of tables ..................................................................................................................... vi Abstract ........................................................................................................................... vii Chapter 1. Introduction ............................................................................................................ 1 2. History of bryozoan research in Ireland ............................................................... 21 3. Distributional patterns of Bryozoa in Ireland Past and present ........................... 38 4. A taxonomic atlas of the bryozoans or Ireland .................................................... 74 5. Invasive and non-native Bryozoa in Ireland………………………………..…106 6. Bryozoans as epiphytes ...................................................................................... 139 Acknowledgements I would like to express special thanks and appreciation to my advisor Professor Patrick Wyse Jackson, you have been a wonderful advisor to me. I would like to thank you for all your encouragement, patience and time when I needed it most. Your advice and knowledge have been invaluable to my research. I would also like to thank Mary Spencer Jones (NHMUK) for your advice on SEM imaging of Bryozoa and Nigel Monaghan (NHMI) for his contribution of photographs from The National Museum of Ireland and for allowing me access to the Bryozoa collection. I would also like to thank Captain Simon Coate, Harbour Master at Dun Laoghaire Harbour and Mr. David Knott, Safety and environmental manager at Belfast Harbour for allowing me access to your harbours and any support I needed while on site. This research was funded by a Trinity College Studentship grant. Trinity College were also kind enough to provide funding for part of the field work required with a Trinity College Travel grant. Additional funding was provided by a SYNTHESYS grant to enable me to examine specimens at NHMUK and use the SEM equipment. A special thanks to my family. Words cannot express how grateful I am to my Husband for the sacrifices you made on my behalf and for all the cups of tea. Your enduring support and love will forever be remembered. I would also like to thank my friends for your words of encouragement at just the right times. iv List of Figures Chapter1 Fig. 1.1. Classification Tree of Bryozoa. Fig. 1.2 SEM image of a calcareous branch section of Cellaria fistulosa from Strangford Lough, Down. Chapter 2 Fig. 2.1 Timeline of bryozoan research in Ireland Fig. 2.2. Albert Russell Nichols (1859-1933) at opening of the Museum of Science and Art, Kildare Street building in 1890. (Photograph NMINH- PP-777xy © National Museum of Ireland, all rights reserved) Fig. 2.3. 30 Grosvenor Square, Rathmines, Dublin (Photograph by Prof. Patrick Wyse Jackson) Fig. 2.4. A.R. Nichols towards the end of his working life. (Photograph NMINH-PP-741-Nichols © National Museum of Ireland, all rights reserved) Fig. 2.5. Burial place of A.R. Nichols, his wife and daughter. Mount Jerome Cemetery, Harold’s Cross, Dublin. (Photograph by Prof. Patrick Wyse Jackson) Fig. 2.6. Cover of Nichols’ contribution on bryozoans to the Clare Island Survey (1912) Chapter 3 Fig. 3.1. North and south divide of Ireland used in this study Fig. 3.2. Settlement panel in Belfast harbour. A) The marker where the panel is attached. B) The settlement panel before placement. C) The panel frame being lowered from the wooden platform. D) Map of Belfast Harbour indicating the location of the panel frame. Fig 3.3. Locations sampled during the sampling programme in the current research from 2011-2015. v Fig 3.4. Location of all bryozoan records taken from the historical and museum records. Fig 3.5. Comparison of Bryozoa species richness in Ireland between 1850-2017 Chapter 4 Fig 4.1. Scruparia chelata (Linnaeus, 1758) Fig. 4.2. Pyripora catenularia (Fleming, 1828) Fig. 4.3. Electra pilosa (Linnaeus, 1767) Fig. 4.4. Flustra foliacea (Linneaus, 1758) Fig. 4.5. Callopora dumerilii (Audouin, 1826) Fig. 4.6. Alderina imbellis (Hincks, 1860) Fig. 4.7. Tricellaria inopinata d’Hondt & Occhipinti Ambrogi, 1985 Fig. 4.8. Cribrilina punctata (Hassall, 1841) Fig. 4.9. Collarina balzaci (Audouin, 1826) Fig. 4.10. Cribrilaria innominata (Couch, 1844) Fig. 4.11. Figularia figularis (Johnston, 1847) Fig 4.12. Chorizopora brongniartii (Audouin, 1826) Fig. 4.13. Escharella immersa (Fleming, 1828) Fig. 4.14. Escharella variolosa (Johnston, 1838) Fig. 4.15. Neolagenipora collaris (Norman, 1867) Fig. 4.16. Neolagenipora eximia (Hincks, 1860) Fig. 4.17. Porella concinna (Busk, 1854) Fig. 4.18. Cryptosula pallasiana (Moll, 1803) Fig 4.19. Schizoporella unicornis (Johnston in Wood, 1844) Fig. 4.20. Escharina johnstoni (Quelch, 1884) Fig. 4.21. Phaeostachys spinifera (Johnston, 1847) vi Fig. 4.22. Fenestrulina malusii (Audouin, 1826) Fig.4.23. Prenantia cheilostoma (Manzoni, 1869) Fig. 4.24. Watersipora subatra (Ortmann, 1890) Fig. 4.25. Turbicellepora avicularis (Hincks, 1860) Fig. 4.26. Disporella hispida (Fleming, 1828) Chapter 5 Fig. 5.1. The invasive non-native species Watersipora subatra from Dun Laoghaire Harbour, Dublin. (M11 Authors collection). Fig. 5.2. Tricellaria inopinata, frontal view showing the internode, ovicells, spines, avicularium and scutum, from Dun Laoghaire Harbour, Dublin. (M3 Authors collection). Chapter 6 Fig. 6.1. Epiphytic Bryozoa. A) Specimen D53 Authors collection: Hydroid encrusted with Crisia sp., a cyclostome and Electra pilosa, collected from Portmarnock Beach, Dublin B) Specimen MH07 Authors collection: Red algae encrusted with E. pilosa collected from Bettystown beach, Meath C) Specimen MH05 Authors collection: Flustra foliacea with Bugulina sp. as an epiphyte collected from Bettytown beach, Meath. D) Specimen D43 Authors collection: Lamininaria sp. with Membranipora membranacea growing on the stipe collected from Balbriggan, Dublin. E) Specimen MH04 Authors collection: Brown algae with multiple colonies of E. pilosa, M. membranacea, Bugulina sp. and Cryptosula pallasiana collected from Bettystown beach, Meath. Fig. 6.2. Specimen H22 Weyman (1997) collection: Electra pilosa on the red algae Porphyridium purpureum collected from Howth, Dublin. Fig. 6.3. Percentage of Bryozoa collected as epiphytes on Irish coast. Fig. 6.4. Substrate utilization by epiphytic bryozoans from the coast of Ireland. vii List of Tables Chapter 1 Table 1.1. Distinguishing features between Ctenostomatida and Cheilostomatida. (Data from Ryland 1970). Chapter 3 Table 3.1. Summary of bathymetric distribution based on zooid type from Stach (1936) Table 3.3. Summary of Irish marine bryozoan’s taxic database Table 3.3. Species found in the highest percentage of counties from both the historical and recent records. Table 3.4. Species richness for North Ireland East coast (NEI)*, Central Ireland East coast (CEI)*, South Ireland East coast (SEI)* over the time periods 1800- 1900 Table 3.5. Type and figured specimens from Ireland held at NHM London Table 3.6 List of bryozoan species collected during the sampling programme of the current study from the coasts of Ireland with notes on location and substrate. Chapter 5 Table 5.1. Species identified during sampling programme of current study and the locations they were recorded at. Chapter 6 Table 6.1. Location bryozoans were collected from during the sampling programme of the current study. viii Abstract Bryozoans are sessile filter feeding marine invertebrates. Research in Ireland on bryozoans has been sparse in recent years but many studies were carried out in the early 1900’s by researchers such as Albert Russell Nichols (1859-1933). This study has a number of interlinked aims. It will evaluate the history of bryozoan research in Ireland and discuss the biologist Albert Russell Nichols (1859-1933) and his contribution to bryozoan research. Bryozoan diversity and distributional changes from Nichols’ time have been examined very little, this study will examine any changes exhibited over the last 150 years. These changes will be evaluated using both a north south and east west divide by looking at historical records and the results of a recent sampling programme. A taxonomically ordered atlas using SEM images and photographs has been compiled of the bryozoans recorded from Ireland using both recent and historical records. In the last 15 years 4 non-native bryozoan species have been recorded in Ireland. Watersipora subatra, Tricellaria inopinata, Bugula neritina and Schizoporella japonica and other potential invasive bryozoans are discussed to evaluate their