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Trophic Ecology in Marine Ecosystems from the Balearic Sea (Western Mediterranean)

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Trophic Ecology in Marine Ecosystems from the Balearic Sea (Western Mediterranean) TROPHIC ECOLOGY IN MARINE ECOSYSTEMS FROM THE BALEARIC SEA (WESTERN MEDITERRANEAN) MARIA VALLS MIR PhD THESIS 2017 I II DOCTORAL THESIS 2017 Doctoral Program of Marine Ecology TROPHIC ECOLOGY IN MARINE ECOSYSTEMS FROM THE BALEARIC SEA (WESTERN MEDITERRANEAN) Maria Valls Mir Thesis Supervisor: Antoni Quetglas Thesis tutor: Gabriel Moyà Doctor by the Universitat de les Illes Balears III IV TABLE OF CONTENTS Agradecimientos VI List of papers VIII List of acronyms and abreviations IX Summary/Resum/Resumen XI Chapter 1. Introduction 1 1.1 Thesis motivation 3 1.2 Bentho-pelagic coupling 4 1.3 Food webs as a basis for an ecosystem based management 5 1.4 The study area: the Balearic Sea 6 1.5 Marine food webs from the Balearic Islands 8 1.6 Trophic studies in the Balearic Sea 9 1.7 Study species 11 1.7.1 Elasmobranchs 11 1.7.2 Cephalopods 12 1.7.3 Mesopelagic fishes 13 1.8 Methodological approaches 14 1.9 Aims 16 Chapter 2. Material and methods 17 2.1 Datasets 19 2.1.1 Scientific surveys 19 2.1.1.1 MEDITS program 19 2.1.1.2 IDEADOS project 20 2.1.1.3 Data Collection Framework (DCF) 21 2.2 Sampling 22 2.2.1 Stomach contents analysis 22 2.2.2 Stable isotope analysis 24 2.2.2.1Lipid content 27 Chapter 3. Structure and dynamics of food webs in the water column on shelf and slope grounds of the western Mediterranean 31 3.1. Introduction 33 3.2 Material and methods 34 3.3 Results 41 3.4 Discussion 45 V Chapter 4. Feeding ecology of demersal elasmobranchs from the shelf and slope off the Balearic Sea (western Mediterranean) 53 4.1 Introduction 55 4.2 Material and methods 55 4.3 Results 56 4.4 Discussion 60 Chapter 5. Feeding ecology of two squid species from the western Mediterranean 5.1 Introduction 71 5.2 Material and methods 72 5.3 Results 74 5.4 Discussion 81 Chapter 6. Trophic structure of mesopelagic fishes in the western Mediterranean based on stable isotopes of carbon and nitrogen 91 6.1 Introduction 93 6.2 Material and methods 95 6.3 Results 98 6.4 Discussion 105 Chapter 7. Feeding strategies and resource partitioning among elasmobranchs and cephalopods in Mediterranean deep-sea ecosystems 111 7.1 Introduction 113 7.2 Material and methods 115 7.2.1 Data source and study area 115 7.2.2 Diet analyses 115 7.3 Results 121 7.4 Discussion 131 Chapter 8. General Discussion 137 Chapter 9. Conclusions 145 References 151 VI Esta tesis te la dedico a ti mamá. Tú me rodeaste de mar y confiaste en mí. VII VIII Agradecimientos Llegué al Centro Oceanográfico de Baleares-Instituto Español de Oceanografía (COB-IEO) en diciembre de 2003. Me presenté para trabajar de observadora a bordo con la flota de arrastre de Mallorca y los artesanales que faenaban en Cabrera. Había soñado con hacer este trabajo y me pareció una oportunidad de oro. Mi primer embarque fue con El Montgó, un arrastrero de Alcudia. Fue un día de muy mal tiempo y ¡lo pasé fatal! Pero seguí adelante y estuve dos años embarcándome. A pesar de los madrugones, las largas jornadas de pesca y un sueldo justito, aprendí un montón, pude participar en mis primeras campañas oceanográficas y comí mucho pescado. Mis primeras GRACIAS son para los pescadores, por dejarme embarcar con ellos y tratarme siempre con mucho cariño. En Cabrera conocí a Álvaro, el padre de mi hija Mar, una historia especial y muy bonita! En el año 2006 conseguí un contrato del Programa Nacional de Datos Básicos del IEO. Y en el 2008 me saqué una oposición para Ayudante de Investigación y pude incorporarme en el COB con una plaza fija! Ya han pasado 10 años desde que me apunté al doctorado. Fue en el 2007, cuando iniciamos en Baleares las campañas Medits, y yo comencé con los análisis de contenidos estomacales e isótopos. Enric Massutí me ofreció la posibilidad de empezar estos muestreos. GRACIAS Enric, por confiar en mí desde que empecé a trabajar en el COB de muestreadora. Él me dio la oportunidad de aprender y luego organizar y desarrollar los análisis de trofismo en las campañas y los trabajos relacionados que han ido surgiendo desde entonces. También quiero dar las GRACIAS a Toni Quetglas, por aceptar dirigir esta tesis de largo recorrido, corregir mi trabajo siempre a conciencia y abrazarme cuando lo he necesitado. Con él empecé los análisis de dieta de cefalópodos, los cuales requieren de un trabajo minucioso y laborioso. Su cuaderno de dibujos y su colección de referencia de picos de cefalópodos me han sido muy útiles. Además de mi director ha sido mi compañero de “camarote” durante esta travesía. Participé en varias campañas (Demersales, Arsa) para coger rodaje y aprender de los más expertos. Así que mil GRACIAS para todos los que me han enseñado el oficio, especialmente a José Luis Zabala, con el que empecé esta andadura. Y por supuesto a Marian Torres, mi compi M&M durante los primeros años de trofismo en campaña, con la que los intensos días de muestreo e informatización de datos se hacían muy amenos, gracias por tu salero chiquilla! Además quiero agradecer a todos los especialistas que me han ayudado a identificar las presas que han ido apareciendo en los estómagos, muchas veces deformadas y semi digeridas, y sólo reconocibles por pequeños detalles y ojos expertos. En este grupo de asesores están Juan Junoy, Alberto Serrano, Inma Frutos, Joan Cartes, Pere Abelló, Xisco Ordines, Enric Massutí, Beatriz Guijarro, Toni Quetglas, Carmen Barberà, Hilmar Hinz, José Rueda, Montse Ramón, Toni Lombarte, Ana Morillas y Maite Vázquez. Por supuesto quiero dejar constancia de la ayuda de los numerosos IX apuntadores/as en las campañas, que me han ayudado a aligerar la cola de “clientes” durante las largas jornadas a bordo. El mundo de los isótopos estables fue otro camino. A pesar de empezar la recogida de muestras a la vez que los estómagos, no empecé a trabajar los datos hasta un tiempo más tarde. Para empezar quiero darle las GRACIAS a Emil Olafsson, por enseñarme el protocolo de procesado de las muestras y facilitarme el material necesario para ello. En el procesado de muestras han hecho mucho trabajo Mª Teresa Farriols, Nuria Zaragoza, Balbina Molí, Carmen Barberà y Pilar Bennasar, GRACIAS a todas! Por supuesto, un montón de GRACIAS para Miquel Ribas y Biel Martorell de la UIB, por enseñarme a trabajar con el espectrofotómetro de masas y estar disponibles siempre (incluido los días festivos) en todas las incidencias que ha tenido la máquina, que han sido muchas! También quiero agradecer a Nick Polunin la oportunidad de trabajar en Newcastle con su equipo y en especial a Christopher Sweeting, por su enseñanza en la interpretación de los datos isotópicos y su inmensa paciencia con todas mis dudas. Gracias también a Pere Oliver por proporcionarme las referencias sobre la historia “trófica” de estas islas. X List of papers This thesis is based on the following papers: I. M. Valls, A. Quetglas, F. Ordines, J. Moranta (2011). Feeding ecology of demersal elasmobranchs from the shelf and slope off the Balearic Sea (western Mediterranean). Scientia Marina 75 (4), 633-639 Impact factor (2015-16): 1.19 II. M. Valls, C.J. Sweeting, M.P. Olivar, M.L. Fernández de Puelles, C. Pasqual, N. Polunin, A. Quetglas (2014). Structure and dynamics of food webs in the water column on shelf and slope grounds of the western Mediterranean. Journal of Marine Systems 138, 171–181 Impact factor (2015-16): 2.174 III. M. Valls, M.P. Olivar, M.L.F. de Puelles, B. Molí, A. Bernal, C.J. Sweeting (2014). Trophic structure of mesopelagic fishes in the western Mediterranean based on stable isotopes of carbon and nitrogen. Journal of Marine Systems 138, 160–170 Impact factor (2015-16): 2.174 IV. M. Valls, M. Cabanellas-Reboredo, I. Uranga, A. Quetglas (2015). Feeding ecology of two squid species from the western Mediterranean. Marine Ecology Progress Series 531, 207-219 Impact factor (2015-16): 2.361 V. M. Valls, L. Rueda, A. Quetglas (2017). Feeding strategies and resource partitioning among elasmobranchs and cephalopods in Mediterranean deep-sea ecosystems. Deep Sea Research Part I, Accepted. Impact factor (2015-16): 2.684 XI XII Acronyms and abbreviations A (%) Appareance frequency ANOVA Analysis of Variance AsB Algerian sub-Basin B Biomass BBL Benthopelagic Boundary Layer Bi Levin’s standardized index BPC Bentho-Pelagic Coupling BsB Balearic sub-Basin C:N Carbon Nitrogen ratio CA/C Cabrera CD Centroid Distance CS Coastal Shelf DCF Data Collection Framework DR Depth Range DSL Deep Scattering Layer GSA Geographical Sub-Area H’ Trophic diversity HC Hyperbenthic Compartment IDEADOS Influencia de la estructura y dinámica oceanográfica sobre poblaciones demersales en aguas de las Islas Baleares IKMT Isaacs-Kidd Midwater Trawl Net IRI Index of Relative Importance MC Monte Carlo test MDS Multidimensional Scaling MEDITS Mediterranean International Trawl Survey ML Dorsal Mantle Length MOCNESS Multiple Opening/Closing Net and Environmental Sensing System MS Middle Slope MSFD Marine Strategy Framework Directive N (%) Percentage by Number of the food item N Total number of individuals O (%) Frequency of occurrence of the food item PC Pelagic Compartment pCCA partial Canonical Correspondence Analysis PERMANOVA Permutational Multivariate Analysis of Variance POM Particulate Organic Matter R/V Research Vessel Reg Regurgitated RMT Rectangular Midwater Trawl SB Shelf Break SCA Stomach Content Analysis SEA Standard Ellipse Area SI Schoener Index SIA Stable Isotope Analysis SIAR Stable Isotope Analysis in R SIMPER Similarity Percentage SIMPROF Similarity Profile analysis XIII SL Standard Length SO Sóller TA Total convex hull Area TDF Trophic discrimination factor TL Total Length Tmax Maximum size Tmin Minimum size TrG Trophic Group TrL Trophic Level TW Total Weight US Upper Slope V (%) Percentage by Volume of the food item v (%) Vacuity index V.E Variance Explained VG Vertical Guild WR Weight Range Isotopic discrimination δ Delta notation Δ XIV Summary Conservation of marine ecosystem structure and functioning is a priority target within the context of environmental management.
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