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POPULATION DYNAMICS OF A NON-INDIGENOUS COLONIAL ASCIDIAN TUNICATE IN A SUBARCTIC HARBOUR by ©KEVIN C. K. MA A thesis submitted to the School of Graduate Studies in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE DEPARTMENT OF BIOLOGY, FACULTY OF SCIENCE MEMORIAL UNIVERSITY OF NEWFOUNDLAND December 2012 ST. JOHN'S NEWFOUNDLAND AND LABRADOR CANADA ABSTRACT Botryllus schlosseri (Subphylum Tunicata: Class Ascidiacea) is a non indigenous asci dian species of global and national interest, which has extensive populations along the south coast of insular Newfoundland. Economically, this species has been of concern to industry, management, and policymakers because non-indigenous ascidian species have been a severe and costly nuisance for bivalve aquaculture. Ecologically, the presence of this temperate-adapted species in Newfoundland represents an expansion of its global range into subarctic waters. Thus, I aimed to describe the population dynamics of B. schlosseri in Arnold's Cove, Placentia Bay, Newfoundland and Labrador, Canada, by determining the temporal and spatial patterns of recruitment and the seasonal cycle of colony abundance. In addition, I aimed to compile a checklist of extant indigenous and non-indigenous ascidian species of eastern Canada with an emphasis on species from Newfoundland and Labrador. Artificial plates were used to determine recruitment rates among three sites, depths (1.0, 2.5, and 4.0 m from the water surface), and substrate types (aluminum, PVC, and wood in 2010; only PVC in 2011), in Arnold's Cove. Concurrently, density and cover of colonies were determined from the analysis of high-definition video surveys of a belt transect of wharf pilings. Seasonal biomass production was estimated from carbon to nitrogen (C:N) ratios and dry weight per unit area of dissected tissue subsamples. ii The seasonal window for recruitment was from early August to mid-October. Recruitment rates were greater near the water surface than at other depths, and on PVC in comparison to aluminum and wood substrates. Maximum recruitment rates on PVC at 1.0 m were 29.3 and 43.5 m-2d-1 in September of 2010 and 2011, respectively, coincident with maximum seasonal seawater temperatures of 16-17°C. Colonies were present year-round on pilings. In the upper subtidal zone, monthly mean cover ranged from an annual minimum of 0.6% in May to a maximum of 2.8% in October. Colony size and biomass, though not C:N ratios, had a significant seasonal signal. These findings suggest that recruitment was predominantly constrained by seawater temperature within the short productive season, and that the population was sustained from one year to the next because of high cover of overwintering colonies. The efficacy of utilising PVC to track recruitment of Botryllus schlosseri, and perhaps other closely related ascidian species is supported by my data. Future management of B. schlosseri should target mitigation efforts before the annual onset of sexual reproduction and recruitment in July and within the upper 3-4 m of the water column. iii TABLE OF CONTENTS Abstract. ...................................................................................................................................................... ii List of tables ............................................................................................................................................. vi List of figures ........................................................................................................................................... xi List ofplates ........................................................................................................................................... xvi Acknowledgments ............................................................................................................................ xviii Co-authorship statement ................................................................................................................... xx Abbreviations ........................................................................................................................................ xxi Chapter 1 - Introduction and literature review ......................................................................... 1 1.1 Introduction and objectives ................................................................................... 1 1.2 Non-indigenous ascidian species ......................................................................... 3 1.3 The biology of Botryllus schlosseri ....................................................................... 9 Chapter 2 -An account of extant ascidians of eastern Canada ........................................... 18 2.1 Introduction ............................................................................................................... 18 2.2 Materials and methods ........................................................................................... 20 2.3 Results .......................................................................................................................... 27 2.4 Discussion ................................................................................................................... 36 Chapter 3 - Patterns of recruitment in a population of Botryllus schlosseri.. ................ 40 3.1 Introduction ............................................................................................................... 40 3.2 Materials and methods ........................................................................................... 42 3.3 Results .......................................................................................................................... 51 3.4 Discussion ................................................................................................................... 69 Chapter 4- Patterns of abundance of Botryllus schlosseri on wharf pilings .................. 79 4.1 Introduction ............................................................................................................... 79 4.2 Materials and methods ........................................................................................... 80 4.3 Results .......................................................................................................................... 89 4.4 Discussion ................................................................................................................ 103 iv Chapter 5 -Conclusion .................................................................................................................... 108 5.1 Summary...... ............................................................................................................. 108 5.2 Discussion ................................................................................................................ 111 5.3 Conclusion ................................................................................................................ 116 Appendix A- Supplemental tables and figures ...................................................................... 118 A-1 Supplemental tables ............................................................................................. 118 A-2 Supplemental figures ........................................................................................... 132 Appendix B-Methods for modeling zero-inflated continuous ecological data ........ 138 Abstract ................................................................................................................................... 138 B-1 Introduction ............................................................................................................ 139 B-2 Methods and results ............................................................................................. 141 B-3 Discussion ................................................................................................................ 149 Appendix C- The oozooid and the first blastozooid of Botryllus schlosseri ............... 150 Abstract ..................................... .............................................................................................. 150 C-1 Introduction ............................................................................................................ 151 C-2 Materials and methods ........................................................................................ 152 C-3 Results ....................................................................................................................... 154 C-4 Discussion ................................................................................................................ 161 Appendix D- Dichotomous key to morphotypes of Botryllus schlosseri ...................... 163 Abstract ................................................................................................................................... 163 D-1 Introduction ............................................................................................................ 163 D-2 Methods and results ............................................................................................. 169 D-3 Discussion ......................................................................... ....................................... 184 Bibliography ........................................................................................................................................ 186 v LIST