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Risk Taking in Semibalanus Balanoides

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Risk Taking in Semibalanus Balanoides Risk taking in Semibalanus balanoides By Amanda Tomas A thesis submitted for the degree of Doctor of Philosophy School of Biology Newcastle University. August 2014 Abstract This thesis examined factors influencing the success of the larval cyprid phase, and the distribution of adult Semibalanus balanoides in the Clyde Sea. Experiments were conducted in the Clyde Sea area (2004 – 2007) because it is effectively both a closed system in its whole extent for release of nauplii and cypris settlement, and at a larger scale it is an open system for the actual settlement on individual rock types. The effect of geology, exposure and position on the shore is examined in relation to barnacle fecundity (number of eggs produced) with metamorphic rock, exposed shores and mid shore settlement of Semibalanus balanoides having higher barnacle fecundity than igneous and sedimentary rock types, sheltered shores and upper or lower shore height areas. Frustum volume was found to be a reliable non-destructive way to estimate fecundity, and egg size did not alter significantly from year to year. Supply was most reliably measured using pumps or adhesive panels, and artificial rock panels as an alternative to natural rock; the majority of cypris left after four seconds investigating artificial panels. Cannibalism takes place in the Clyde Sea with 44 % of adults investigated on the shore showing signs of this, frustum size and height didn’t affect propensity to cannibalise. Video recording demonstrated a mean of nine cypris and / or nauplii consumed per adult per hour. Settlement occurred on all major rock types in the Clyde Sea. Cypris preferentially settled on metamorphic and igneous rock types. There was no significant effect of north or south orientation on vertical panels for settlement intensification. Settlement levels were higher on natural rock substratum than artificial panels, but were not significantly different; and showed larger per area settlement on smaller areas as 1 cm2 had a mean settlement of 49.40 cypris, 5 cm2 18.40, 10 cm2 22.42, 50 cm2 13.92, 100 cm2 9.34 and 500 cm2 5.65. On natural rock around the Clyde Sea (as opposed to areas cleared for experiments), areas of < 1 cm2 and 1 cm2 are more common as potential settlement sites, comprising of 51.99 % and 23.98 %, respectively of available settlement area. The mean number of barnacles in the Clyde Sea area was calculated to be 9.454 x 1011, which could produce a mean number of 1.444 x 1015 eggs. i Acknowledgements I would like to thank my initial supervisor, Dr Jeremy Thomason for his constant support even when he left the university. He encouraged, inspired and advised me throughout. His patience in teaching statistics was truly amazing as was his broad knowledge of the marine environment. Dr Gordon Port was extremely generous with his time, advice and patience as first of all my second supervisor then my only supervisor; I couldn’t have had better supervisors. This thesis would not have been possible without the support of my husband Neil who believed in me and advised me on the mapping; the patience, fortitude and understanding of my daughter Cara, son Adam and Jessica the cat (who would cuddle into me). The School of Biology was incredibly supportive and understanding of my illness (ME) beyond the call of duty, and I really appreciated it. The work would not have been possible without the assistance in field work, Dr Phil Cowie deploying a frame from the pier at the University Biological Station, Millport and collecting adult Semibalanus balanoides when I was unable to get to Cumbrae. Colleagues in the Research Group helped in both field work and advice especially H. E. Sugden, G. S. Prendergast, H. Naser and C. M. Zurn. I would also like to thank the staff at the University Biological Station, and Fairlie Pier who both allowed access and experiments to be deployed from their piers. I would like to thank the examiners Dr Phil Cowie (External) and Professor Matt Bentley (Internal) who provided much helpful advice during my viva. ii Contents 1 Introduction 1 1.1 Characteristics of rocky shores and threats to rocky shore communities 1 1.2 Barnacles fouling boats and installations such as power stations 7 1.2.1 Barnacle adhesion 8 1.3 Semibalanus balanoides 10 1.3.1 Why choose this species? 10 1.3.2 Life history and Morphology 11 1.3.3 Distribution and abiotic factors 16 1.3.4 Supply side ecology 17 1.3.5 Predators 21 1.3.6 Gregariousness 22 1.3.7 Population (intraspecific competition) 22 1.3.8 Community (interspecific competition) 23 1.3.9 Post settlement mortality 24 1.3.10 Genetic effects 24 1.4 Why choose the Clyde Sea area? 25 1.5 Overview of thesis 27 1.5.1 Aims and objectives of thesis 27 2 Mapping, geology, bathymetric data and development of exposure ratings 30 2.1 Introduction 30 2.1.1 Clyde Sea area 30 2.2 Aims of chapter 33 2.3 Methods and materials 34 2.3.1 Mapping of the foreshore and its features 34 2.3.2 Geological mapping 36 2.3.3 Bathymetric mapping 39 2.3.4 Development of shore exposure ratings 46 2.3.5 Data analysis 48 2.4 Results 48 2.4.1 Mapping of the foreshore and its features 48 2.4.2 Geological mapping 54 2.4.3 Bathymetric mapping 57 2.4.4 Combined mapping to OSGB 36 59 iii Contents 2.4.5 Exposure rating of different shores 61 2.5 Discussion 68 2.6 Conclusions 72 3 Fecundity 73 3.1 Introduction 73 3.2 Aims of chapter 79 3.3 Methods and materials 80 3.3.1 Sample collection 80 3.3.2 Non-destructive estimation of fecundity 85 3.3.3 Determination of fecundity 87 3.3.4 Data analysis 89 3.4 Results 90 3.4.1 Fecundity estimates 90 3.4.2 Relationship between rock type, exposure, density, shore height and barnacle density 97 3.4.3 Fecundity, egg size and somatic tissue for 2004 and 2005, fecundity and egg size for 2006 105 3.5 Discussion 109 3.6 Conclusions 115 4 Cyprid supply and the influence of substratum 116 4.1 Introduction 116 4.2 Aims of chapter 124 4.3 Methods and materials 125 4.3.1 Cyprid supply and settlement frame 125 4.3.2 Pumps 125 4.3.3 Traps 128 4.3.4 Plain settlement panels 128 4.3.5 Adhesive settlement panels 128 4.3.6 Deployment 128 4.3.7 Filming of cypris arrival, departure and settlement on artificial rock panels 129 4.3.8 Cypris settlement on natural rock versus artificial rock panels 131 4.3.9 Data recording and analysis 132 iv Contents 4.4 Results 134 4.4.1 Cyprid supply and settlement frame 134 4.4.2 Filmed settlement on artificial rock panels 139 4.4.3 Natural versus artificial rock panels 143 4.5 Discussion 144 4.6 Conclusions 149 5 Cannibalism 150 5.1 Introduction 150 5.2 Aims of chapter 153 5.3 Methods and materials 153 5.3.1 Gut contents analysis 154 5.3.2 Filming of cannibalism 154 5.3.3 Data analysis 155 5.4 Results 157 5.4.1 Is Semibalanus balanoides cannibalistic? 157 5.4.2 Does the size of the adult affect cannibalistic tendancies 160 5.4.3 Encounter and capture rates 161 5.4.4 Is increased height of adult an indicator to cannibalism? 163 5.4.5 Does state of the tide affect cannibalism? 164 5.5 Discussion 167 5.6 Conclusions 172 6 Effect of rock type on settlement 173 6.1 Introduction 173 6.2 Aims of chapter 178 6.3 Methods and materials 179 6.3.1 Data analysis 187 6.4 Results 187 6.4.1 Effect of rock type on settlement 187 6.5 Discussion 189 6.6 Conclusions 192 7 Effect of available substratum area on settlement intensification 193 7.1 Introduction 193 7.2 Aims of chapter 195 v Contents 7.3 Methods and materials 196 7.3.1 Settlement on artificial panels 196 7.3.2 Settlement on natural (rock) substratum 198 7.3.3 Space available for settlement 202 7.3.4 Data analysis 202 7.4 Results 203 7.4.1 Settlement on artificial panels 203 7.4.2 Settlement on natural (rock) substratum 205 7.4.3 Space available for settlement 209 7.5 Discussion 210 7.6 Conclusions 214 8 Final Discussion 215 9 References 230 vi Contents List of figures Figure 1-1 Semibalanus balanoides adults, newly metamorphosed juveniles and cypris 11 Figure 1-2 Configuration of the cirri (numbers 1-6 which are altered thoracic limbs) of Semibalanus balanoides. 13 Figure 1-3 Drawing of the United Kingdom with the Clyde Sea Area in the small box 26 Figure 1-4 Diagram of thesis overview and how chapters are linked together 29 Figure 2-1 Illustration of Gnomonic projection 40 Figure 2-2 Ordnance Survey data tidied 49 Figure 2-3 Mean Low Water Springs (MLWS in green) and Mean High Water Springs (MHWS in red) of the Clyde Sea area 51 Figure 2-4 MHWS with places shown for data collection and experiments in Clyde Sea area 53 Figure 2-5 Geology of the Clyde Sea area with MHWS to show the foreshore 55 Figure 2-6 Bathymetric map of Clyde Sea area, of all charts to OS36 projection and MHWS and MLWS 58 Figure 2-7 Combined map of Clyde Sea area, of Ordnance Survey data, geology and bathymetric data (OS36 projection) 60 Figure 2-8 Rose diagram or compass rose divided into 16 segments used to estimate fetch.
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