Population structure of the bluemouth, Helicolenus dactylopterus (Teleostei: Sebastidae), in the Northeast Atlantic and Mediterranean using geometric morphometric techniques Rebeca Paola Rodríguez Mendoza PhD Thesis Vigo, 2013 Depto. de Ecología y Biología Animal Facultad de Ciencias Population structure of the bluemouth, Helicolenus dactylopterus (Teleostei: Sebastidae), in the Northeast Atlantic and Mediterranean using geometric morphometric techniques Memoria de Tesis Doctoral para optar al grado de Doctor por la Universidad de Vigo Presentada por: Rebeca Paola Rodríguez Mendoza Vigo, 2013 A mis padres, a mis hermanos, a Alejandro, a toda mi familia, y a mis amigos… Acknowledgments | Agradecimientos Quiero agradecer a mis padres, por haberme apoyado en todos los aspectos de mi vida y haberme animado a que siguiera adelante en todo momento, a pesar de la distancia que nos separa. A mis hermanos, a Margarita y Álvaro por su apoyo y su cariño. A mi pequeña Sofi que me ha traido tantas alegrías, a Ivancito y a Emil por darme tantas ilusiones y fuerzas para seguir adelante. También a Alejandro, junto con quien he formado una familia y me da su amor y cariño todos los días. A sus padres, que están siempre ahí apoyándonos en todo. ¡Gracias a toda mi gran familia! Quiero agradecer a todos mis amigos, en México, en España y en Croacia ¿qué habría hecho yo en tantas ocasiones sin su apoyo? Gracias queridas amigas: Sonia, Iramaia, Fiona, Consuelo, Loli… Muchas gracias a Mak, sin quien no hubiera sido capaz de llegar hasta aquí. A Jose por todo su apoyo y cariño. Quiero expresar mi sincero agradecimiento a Fran Saborido Rey por darme la oportunidad de entrar en este mundo de la ecología pesquera. ¡Muchas gracias por apoyarme en mi formación! También quiero agradecer a todos mis compañeros del equipo de pesquerías, por su apoyo, emocional y profesional. Asimismo, a todos aquellos que han colaborado directa o indirectamente en esta tesis, ya que gracias a ustedes, este trabajo es lo que es. Posebno se želim zahvaliti Prof. Tomislavu Treeru i Marini Piria, jer su mi puno pomogli da stignem toliko daleko. Special thanks to Dr. Klingenberg, for sharing his knowledge and answering my long e-mails. Thanks to Prof. Rohlf, to Andrea Cardini and all of you who have made accessible to biologists all the theory of geometric morphometrics! Finally, thanks to the ECOSUMMER project, to Graham and Bego, Vasilis and all of my friends! Participating Institutions This PhD Thesis was developed within the PhD Program “Biology and Ecology of Organisms” of the University of Vigo, Spain. Research was carried out at the Institute of Marine Research in Vigo - Spanish Research Council (IIM-CSIC) under the supervision of Dr. Fran Saborido-Rey and at the Spanish Institute of Oceanography in Vigo (IEO) - Ministry of Economy and Competitiveness, under the supervision of Dr. Graham J. Pierce. The author’s work was mainly funded by a Marie Curie fellowship from the Early Stage Training site ECOSUMMER (MEST-CT-2005-020501) and a CONACyT - Mexico scholarship. This research was also supported by the project CTM2006-13964-C03-00 of the Spanish Ministry of Science and Innovation. Table of Contents SUMMARY ................................................................................................................................................ 17 CHAPTER 1 - GENERAL INTRODUCTION ...................................................................................... 23 1.1 OVERVIEW ........................................................................................................................................... 25 1.2. THE BLUEMOUTH ............................................................................................................................. 25 1.2.1 TAXONOMY ............................................................................................................... 25 1.2.2 MORPHOLOGY ............................................................................................................... 28 1.2.3 DISTRIBUTION AND HABITAT ................................................................................................. 28 1.2.4 BEHAVIOUR ............................................................................................................... 32 1.2.5 AGE AND GROWTH ............................................................................................................... 32 1.2.6 DIET ............................................................................................................................. 37 1.2.7 REPRODUCTION ............................................................................................................... 37 1.3 DEEP-SEA FISHERIES ............................................................................................................... 40 1.3.1 OVERVIEW OF DEEP-SEA FISHERIES ................................................................................... 40 1.3.2 IMPACTS OF DEEP-SEA FISHING ................................................................................... 41 1.3.3 ASSESSMENT AND MANAGEMENT OF DEEP-SEA FISHERIES ........................................ 42 1.3.4 CURRENT SITUATION OF THE BLUEMOUTH IN THE FISHERIES ........................................ 44 1.4 POPULATION STRUCTURE OF MARINE FISH AND MANAGEMENT ...................................................... 48 1.4.1 STOCK IDENTIFICATION ................................................................................................. 48 1.4.2 GEOMETRIC MORPHOMETRICS AND THE STUDY OF MARINE POPULATIONS ............ 51 1.5 AIMS AND OBJECTIVES ............................................................................................................... 53 CHAPTER 2 - MATERIAL AND METHODS ..................................................................................... 57 2.1 SPECIMENS AND SAMPLING LOCATIONS ................................................................................... 59 2.2 SELECTION AND PROCESSING OF SAMPLES ................................................................................... 67 2.3 MORPHOMETRICS ............................................................................................................................. 70 2.3.1 STATISTICAL ANALYSIS OF MORPHOMETRIC DATA ...................................................... 73 2. 3. 2 DETECTION OF OUTLIERS, QUANTIFICATION OF MEASUREMENT ERROR AND EVALUATION OF BILATERAL SYMMETRY ............................................................................................................... 76 2. 3. 3 STATISTICAL ANALYSES ....................................................................................................... 81 2.4 MERISTICS ........................................................................................................................................... 93 2.4.1 STATISTICAL ANALYSIS OF MERISTIC VARIABLES .................................................................. 96 Table of Contents (continued) CHAPTER 3 - ONTOGENETIC ALLOMETRY AND SIZE CORRECTION ............................... 101 3.1 INTRODUCTION ........................................................................................................................... 103 3.2 OBJECTIVES ........................................................................................................................... 107 3.3 RESULTS ......................................................................................................................................... 108 3.3.1 CHARACTERIZATION OF ALLOMETRIC SHAPE TRAJECTORIES ....................................... 108 3.3.2 COMPARISON OF THE SIZE-CORRECTION METHODS ..................................................... 122 3.4 DISCUSSION ........................................................................................................................... 128 3.5 CONCLUDING REMARKS ............................................................................................................. 136 CHAPTER 4 - POPULATION STRUCTURE BASED ON GEOMETRIC MORPHOMETRICS AND MERISTICS ................................................................................................................................... 137 4.1 INTRODUCTION ........................................................................................................................... 139 4.1.1 POPULATION STRUCTURING OF MARINE FISH IN THE NE ATLANTIC AND MEDITERRANEAN ......................................................................................................................................... 139 4.1.2 BLUEMOUTH POPULATION STRUCTURE IN THE NE ATLANTIC AND WESTERN MEDITERRANEAN ............................................................................................................. 141 4.2 OBJECTIVES ........................................................................................................................... 144 4.3 RESULTS ......................................................................................................................................... 145 4.3.1 SEXUAL DIMORPHISM ............................................................................................... 145 4.3.2 OVERALL MORPHOMETRIC ANALYSIS ................................................................................. 149 4.3.3 OVERALL MORPHOMETRIC ANALYSIS (EXCLUDING SAMPLES FROM 2006) ........................ 151 4.3.4 MORPHOMETRIC ANALYSIS IN THE MEDITERRANEAN ..................................................... 159