Understanding the biology of two commercially important crustaceans in relation to fisheries and anthropogenic impacts Kevin Scott Submitted for the degree of Doctor of Philosophy Heriot Watt University The School of Energy, Geoscience, Infrastructure and Society (EGIS) July 2019 The copyright in this thesis is owned by the author. Any quotation from the thesis or use of any of the information contained in it must acknowledge this thesis as the source of the quotation or information. ABSTRACT The edible crab, Cancer pagurus and the European lobster, Homarus gammarus, are two commercially and ecologically important species found throughout Western Europe. Despite their importance there remains large knowledge gaps about both species, particularly around the effects of recent anthropogenic marine stressors such as unavoidable electromagnetic field (EMF) emissions from Marine Renewable Energy Devices (MREDs). Given the life history of C. pagurus and H. gammarus there is a high likelihood that they will come in to contact with MREDs and their associated power cables. To fully assess the potential impacts of EMF on C. pagurus and H. gammarus several factors were assessed including (1) behavioural aspects such as attraction/avoidance, activity level, and antennular flicking rate, (2) physiological aspects using commonly utilised crustacean stress markers, and (3) developmental aspects via examination of egg development, egg and larval morphology, and larval locomotory ability. Results were compared to two separate studies conducted to expand the knowledge on baseline stress markers for each species. Results confirm that EMF exposure negatively impacts both species on a behavioural, physiological and developmental level with far reaching implications. Results are discussed collectively and implications for management and future research suggested. i ACKNOWLEDGEMENTS This PhD thesis would not have been possible without the support of a great number of individuals and organisations. I am especially grateful for the financial support from the Nesbitt-Cleland Trust without which the completion of this work would not have been possible. I would also like to thank the trustees of St Abbs Marine Station for constant support beyond anything ever expected. I would like to extend my sincere gratitude to my supervisors Dr Alastair Lyndon and Dr Dan Harries for the constant feedback and support. I would like to thank Dr Alastair Lyndon in particular for the advice given throughout my studies and for acting as my professional mentor throughout my career. I would also like to thank Petra Harsanyi who played an integral role in all of the research contained within this thesis, and for all the extra time and effort she put in. I am extremely grateful to the many volunteers and interns who have passed through the doors of St Abbs Marine Station and helped with this research in a great variety of ways, in particular Guadalupe de la Cruz Ortiz. Special thanks also go to the current staff of St Abbs Marine Station; Adam Houghton, Erica Chapman, and Blair Easton for all their hard work over the last several years and for contributing to this work in many ways. I would like to thank the St Abbs fishermen for providing access to their boats, allowing us to conduct surveys and for helping us deploy equipment from their boats. Special thanks to Philip Rutherford for helping with all manner of research aboard his boat. I would like to extend my gratitude to the Scottish Fishermen’s Federation, and Marine Alliance for Science and Technology Scotland for providing funding for various parts of this research. Special thanks to my examiners, Professor Michel Kaiser and Dr Nia Whiteley for providing valuable feedback. A very special thanks go to my loving wife Carol and my son Arran for providing me with a great escape from work and for always being there for me. Finally, I would like to thank my parents, Carol and Kenny, for keeping me on the right path and encouraging me to never give up. ii ACADEMIC REGISTRY Research Thesis Submission Name: Kevin Scott School: Energy, Geoscience, Infrastructure and Society (EGIS) Version: (i.e. First, Final Degree Sought: PhD Marine Biology Resubmission, Final) Declaration In accordance with the appropriate regulations I hereby submit my thesis and I declare that: 1) The thesis embodies the results of my own work and has been composed by myself 2) Where appropriate, I have made acknowledgement of the work of others 3) The thesis is the correct version for submission and is the same version as any electronic versions submitted*. 4) My thesis for the award referred to, deposited in the Heriot-Watt University Library, should be made available for loan or photocopying and be available via the Institutional Repository, subject to such conditions as the Librarian may require 5) I understand that as a student of the University I am required to abide by the Regulations of the University and to conform to its discipline. 6) I confirm that the thesis has been verified against plagiarism via an approved plagiarism detection application e.g. Turnitin. * Please note that it is the responsibility of the candidate to ensure that the correct version of the thesis is submitted. Signature of Date: Candidate: Submission Submitted By (name in capitals): KEVIN SCOTT Signature of Individual Submitting: Date Submitted: iii For Completion in the Student Service Centre (SSC) Received in the SSC by (name in capitals): Method of Submission (Handed in to SSC; posted through internal/external mail): E-thesis Submitted (mandatory for final theses) Signature: Date: iv PhD Project Outputs Peer-reviewed Scott, K., Harsanyi, P. and Lyndon, A.R., 2018. Understanding the effects of electromagnetic field emissions from Marine Renewable Energy Devices (MREDs) on the commercially important edible crab, Cancer pagurus (L.). Marine Pollution Bulletin, 131, pp.580-588. Scott, K., Harsanyi, P. and Lyndon, A.R., 2018. Baseline measurements of physiological and behavioural stress markers in the commercially important decapod Cancer pagurus (L.). Journal of Experimental Marine Biology and Ecology, 507, pp.1-7. Scott, K., Harsanyi, P. and Lyndon, A.R., 2019. Understanding the effects of electromagnetic field emissions from Marine Renewable Energy Devices (MREDs) on the commercially important edible crab, Cancer pagurus (L.). Frontiers Marine Science Conference Abstract: IMMR'18 International Meeting on Marine Research 2018. doi: 10.3389/conf.FMARS.2018.06.00105 Non peer-reviewed Scott, K., Harsanyi, P. and Lyndon, A, 2018. Understanding the effects of electromagnetic field emissions from Offshore Renewable Energy Devices (OREDs) on the commercially important Edible crab, Cancer pagurus (L.). Bulletin of the Porcupine Marine Natural History Society. Porcupine Marine Natural History Society. Scott, K., 2018. Electromagnetic fields and the invisible threat to seabed species. The Marine Biologist, (11). Conference presentations Scott, K., 2019. Kunsan University Offshore Wind Research Committee. Technical Workshop “Electromagnetic Field (EMF) research in relation to offshore wind farms and aquaculture”, South Korea. v Scott, K., 2018. Marine Alliance for Science and Technology for Scotland (MASTS) Annual Science Meeting. “Understanding the effects of electromagnetic field (EMF) emissions from offshore windfarms on commercially important crustaceans”, United Kingdom. Scott, K., 2018. International Meeting on Marine Research (IMMR). Technical Workshop “Understanding the effects of electromagnetic field (EMF) emissions from Marine Renewable Energy Devices (MREDs) on the commercially important edible crab, Cancer pagurus (L.), Portugal. Scott, K., 2018. Porcupine Marine Natural History Society (PMNHS). “The effects of Offshore Renewable Energy Devices (OREDs) on the commercially important brown crab, Cancer pagurus (L.)”, United Kingdom. Scott, K., 2017. Crustacean Society Mid-Year Meeting + Colloquium Crustacea Decapoda Mediterranea. “Understanding the effects of electromagnetic field emissions from Marine Renewable Energy Devices (MREDs) on the commercially important Edible crab, Cancer pagurus (L.)” , Spain. vi TABLE OF CONTENTS Chapter 1. General Introduction 1.1 Ecology, habitat and distribution of Cancer pagurus 1 1.2 Cancer pagurus fishery and management 3 1.3 Life history of Cancer pagurus 5 1.4 Ecology, habitat and distribution of Homarus gammarus 7 1.5 Homarus gammarus fishery and management 9 1.6 Life history of Homarus gammarus 10 1.7 Renewable energy in the marine environment 11 1.8 Project aims 17 1.9 Thesis structure 18 Chapter 2. Baseline measurements of physiological and behavioural stress markers in the commercially important decapod Cancer pagurus (L.) 2.1 Introduction 20 2.2 Methods and Materials 22 2.3 Results 26 2.4 Discussion 31 Chapter 3. Baseline measurements of physiological and behavioural stress markers in the commercially important decapod Homarus gammarus (L.) 3.1 Introduction 36 3.2 Methods and Materials 38 3.3 Results 42 3.4 Discussion 49 vii Chapter 4. Understanding the effects of electromagnetic field emissions from Marine Renewable Energy Devices (MREDs) on the commercially important edible crab, Cancer pagurus (L.) 4.1 Introduction 54 4.2 Methods and Materials 56 4.3 Results 64 4.4 Discussion 71 Chapter 5. Behavioural and physiological responses in the European lobster, Homarus gammarus (L.), to electromagnetic field emissions from Marine Renewable Energy Devices (MREDs) 5.1 Introduction 77 5.2 Methods and Materials 80 5.3 Results 86 5.4