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Ecology of Carybdea Marsupialis (Cubozoa) and Jellyfish Sting Risk Management

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Ecology of Carybdea Marsupialis (Cubozoa) and Jellyfish Sting Risk Management STUDIES ON THE ECOLOGY OF CARYBDEA MARSUPIALIS (CUBOZOA) AND JELLYFISH STING RISK MANAGEMENT Cesar Bordehore Fontanet STUDIES ON THE ECOLOGY OF CARYBDEA MARSUPIALIS (CUBOZOA) AND JELLYFISH STING RISK MANAGEMENT Thesis submitted by Cesar Bordehore Fontanet for the degree of Doctor of Philosophy in Biology University of Alicante (Spain) September de 2014 ESTUDIOS SOBRE LA ECOLOGÍA DE CARYBDEA MARSUPIALIS (CUBOZOA) Y GESTIÓN DEL RIESGO ASOCIADO A PICADURAS DE MEDUSAS Memoria presentada por Cesar Bordehore Fontanet para optar al título de Doctor en Biología Universidad de Alicante, septiembre 2014 Dissertation Directors / Directores de Tesis: Dra. Verónica L. Fuentes, Instituto de Ciencias del Mar, CSIC, Barcelona. Dr. Alfonso A. Ramos Esplá, Dep. CC Mar y Biología Aplicada, Universidad de Alicante. PhD Tutor / Tutor de Tesis: Dr. Josep Raventós Bonvehí, Dep. Ecología, Universidad de Alicante. Fotografías de portada: Eduardo Obis. Las fotografías de E. Obis han sido realizadas en las instalaciones del Instituto de Ciencias del Mar en Barcelona, CSIC. Dña. Verónica L. Fuentes, Dra en Biología e Investigadora en el Instituto de Ciencias del Mar en Barcelona, del Consejo Superior de Investigaciones Científicas y D. Alfonso A. Ramos Esplá, Catedrático del Departamento de Ciencias del Mar de la Universidad de Alicante, HACEN CONSTAR Que la Tesis Doctoral titulada “STUDIES ON JELLYFISH: ECOLOGY OF CARYBDEA MARSUPIALIS (CUBOZOA) AND JELLYFISH STING RISK MANAGEMENT” recogida en la presente memoria, de la que es autor el Licenciado en Biología D. Cesar Bordehore Fontanet, ha sido realizada bajo nuestra codirección, y cumple las condiciones exigidas para que su autor pueda optar al grado de Doctor por la Universidad de Alicante, por lo que damos la aprobación para su correspondiente lectura y defensa. Para que conste a los efectos oportunos, firmamos el presente escrito el Alicante, a 10 de septiembre de 2014. Dra Verónica L. Fuentes Dr Alfonso A. Ramos Esplá D. Josep Raventós Bonvehí, Catedrático del Departamento Ecología de la Universidad de Alicante, como Tutor de Doctorado del Licenciado en Biología D. Cesar Bordehore Fontanet, Vista la Memoria titulada “STUDIES ON JELLYFISH: ECOLOGY OF CARYBDEA MARSUPIALIS (CUBOZOA) AND JELLYFISH STING RISK MANAGEMENT”, dirigida por la Dra Verónica Fuentes (ICM-CSIC) y el Dr Alfonso A. Ramos Esplá (UA), para optar al grado de Doctor, INFORMA, que reúne las condiciones necesarias para su defensa. Alicante, a 10 de septiembre de 2014. Dr Josep Raventós Bonvehí A Inma, Irene y Lucía, Si Déu ens va posar a la Terra, tenim el dret de fer ús d'ella, però convé que ho fem amb una certa intel·ligència. Ramon Margalef i López (1919-2004) Although we have some knowledge of cubozoan life histories (e.g. Straehler-Pohl & Jarms 2011), detailed descriptions are not available for most taxa. In contrast to their scyphozoan relatives (Arai 1997, Kingsford et al. 2000, Brodeur et al. 2002, Lynam et al. 2006) critical information on temporal and spatial variation in abundance, reproductive output, larval distribution, age, growth, recruitment, mortality and the influence of physical forcing on population dynamics are rare for cubozoans. We have only found two papers that have quantitative data on abundance (Bordehore et al. 2011; Kingsford et al. 2012) [An extract from page 268] Mike J. Kingsford and Christopher J. Mooney (2014) Chapter 12. The Ecology of Box Jellyfishes (Cubozoa) In: Jellyfish Blooms. Eds Kylie A. Pitt and Cathy H. Lucas. Springer ÍNDICE Página AGRADECIMIENTOS 13 CAPÍTULO 1. INTRODUCCIÓN GENERAL 15 1.1. Taxonomía e identificación de cubozoos 16 1.2. Distribución de Carybdea marsupialis 19 1.2.1. Mediterráneo 19 1.2.2. Atlántico, Pacífico e Indico 20 1.3. Ciclo de vida de C. marsupialis 20 1.4. Ecología de cubozoos 23 1.4.1. Distribución espacial y temporal de cubozoos 24 1.4.2 Movimientos y comportamiento 26 1.4.3. Hábitos alimenticios de las cubomedusas 27 1.4.4. Impacto potencial sobre las presas 29 1.4.5. Predadores de cubozoos 29 1.5. Modelos poblacionales de organismos marinos. Aplicación a C. 30 marsupialis 1.6. El problema de las medusas 30 1.6.1. Preocupación pública por las medusas 30 1.6.2. Riesgos en playas 31 1.6.3. Picaduras de medusas 32 1.6.4. C. marsupialis en el área de estudio y salud pública 33 1.6.5. Nematocistos y efectos sobre la salud 34 1.7. Objetivos y estructura de la tesis 35 CAPÍTULO 2. Detection of an unusual presence of the cubozoan Carybdea 37 marsupialis at shallow beaches located near Dénia, Spain (SW Mediterranean) CAPÍTULO 3. Abundance patterns and size structure of Carybdea marsupialis 51 (Cubozoa) at a shallow beach in the West Mediterranean CAPÍTULO 4. Estimating the dynamics and management strategies of marine 69 organisms using an inverse method for time series. An application to the box jellyfish Carybdea marsupialis size structured populations CAPÍTULO 5. Lifeguard assistance at Spanish Mediterranean beaches: 95 Jellyfish prevail and proposals for improving risk management CAPÍTULO 6. Carybdea marsupialis (Cubozoa) in the Mediterranean Sea: 113 The first case of a sting causing cutaneous and systemic manifestations CAPÍTULO 7. DISCUSIÓN GENERAL 121 7.1. Abundancia de C. marsupialis 121 7.2. Factores implicados en la heterogeneidad espacial de la distribución de 124 C. marsupialis 7.3. Ciclo anual 127 7.4. Estrategias reducción de la población de C. marsupialis 127 7.5. Picaduras de medusas y servicio de socorrismo en playas del 128 Mediterráneo español 8. CONCLUSIONES GENERALES 131 GENERAL CONCLUSIONS 133 11 Indice 9. CUESTIONES ABIERTAS Y TRABAJOS FUTUROS 135 9.1. Muestreos de cubozoos 135 9.1.1. Métodos directos 135 9.1.2. Muestreos indirectos 136 9.2. Uso de estatolitos de C. marsupialis 137 9.2.1. Crecimiento 137 9.2.2. Relación Sr/Ca 137 9.3. Localización de los pólipos 138 9.4. Posibles factores ambientales desencadenantes de la metamorfosis 140 9.5. Advección y natación de C. marsupialis 140 9.5.1. Advección 140 9.5.2. Velocidad de natación de diferentes tallas 141 9.6. Distribución de adultos: patrones día-noche y posibles movimientos a 141 gran escala 9.6.1. Posible acercamiento a la costa durante la noche 141 9.6.2. Marcaje de individuos 141 9.7. Reducción de los aportes antrópicos de nitrógeno y fósforo 142 9.8. Datos de picaduras y seguimiento posterior 142 REFERENCIAS 145 ANEXOS I) ACRONYMS / ACRÓNIMOS 159 II) FERTILIZACIÓN DE AGUAS COSTERAS Y REDUCCIÓN DE 161 APORTES DE ORIGEN ANTRÓPICO II.1. Cuantificación de la fertilización de origen antrópico 161 II.2. Propuesta de reducción de la fertilización por N y P de origen 164 antrópico II.2.1. Directiva Nitratos 165 II.2.2. Directiva de aguas subterráneas 166 II.2.3 Directiva aguas residuales urbanas 167 III) ANTECEDENTES Y GÉNESIS PROYECTO LIFE CUBOMED 171 COFINANCIADORES 177 AFILIACIÓN DE COAUTORES 179 12 AGRADECIMIENTOS Quisiera agradecer a todas las personas que han hecho posible la redacción de esta tesis y en especial a: La Dra Verónica Fuentes y al Dr Josep María Gili, ambos del Instituto de Ciencias de Mar de Barcelona (CSIC), por su apoyo e implicación desde el principio de los estudios de cubomedusas en Dénia, allá por el verano de 2008. Sin ellos, ni el proyecto LIFE CUBOMED ni esta tesis habrían visto la luz. Al Dr Alfonso Ramos, por iniciarme en la investigación, allá por el año 1995 –cuando tuve una beca de colaboración del Ministerio de Educación en la antigua Unidad de Biología Marina, hoy Departamento de Ciencias del Mar y Biología Aplicada–, y posterior director de mi beca de FPU y codirector de esta tesis. Ha tenido que esperar casi 14 años más de lo que estaba previsto, un poco de retraso, pero por fin aquí está. Al profesor Josep Raventós, tutor mío de doctorado, y al Dr José Segarra, a ambos por su inestimable colaboración en la modelización de poblaciones y transmitirme su ilusión por hacer nuevas cosas en este apasionante mezcla entre la ecología y las matemáticas aplicadas. A mi compañero de proyecto Dr Just Bayle por todos sus esfuerzos para que el proyecto saliera adelante, sobre todo en los momentos más difíciles. A los investigadores y profesores de las instituciones extranjeras por las que he pasado más o menos tiempo, a Claude y Françoise Monniot, del Museo de Historia Natural de París (Francia, en 1998), a Mike J. Kingsford y Tony Underwood de la Universidad de Sydney (Australia, en 1999) y a Rafael Riosmena de la Universidad Autónoma de Baja California Sur (México, en el año 2000), a todos ellos, quiero agradecer su hospitalidad y formación por unas semanas o meses en sus laboratorios –estancias que, sin saberlo en su momento, me han servido para cumplir uno de los requisitos del Doctorado Internacional–. Agradecer muy especialmente a la Dra Jennifer Purcell, de la Western Washington University (USA) por sus comentarios, correcciones científicas y de lengua inglesa que me han ayudado enormemente a mejorar el contenido de los capítulos 2 al 6. A todos los compañeros que han pasado por el proyecto desde sus inicios: Carmen Barberá, Damián Fernández, Mireia Roig, Elia Durá, Isidro Rico, Antonio Ortiz, Kilian Toledo y Cristina Alonso. También indicar que han sido fundamentales todas las personas que han ido pasando por la Estación Científica Montgó, en períodos de formación de postgrado o realizando sus trabajos de final de grado o máster en Dénia, sin su granito de arena –algunos miles de granitos de arena– habría sido imposible llevar a cabo tantos y tantos muestreos y trabajo de laboratorio, como Beatriz Ramos, Bárbara Jiménez, Aaron Sánchez, Jorge Díez, Aurora Alcover, Arantxa Oquina, Guillermo 13 Agradecimientos Muñoz, Javier García, Lidia Nuño, Lara Saiz, María Gutiérrez, Rebeca Ocón, Leticia Vázquez, Vicente Bernabéu, Neus Figueras, Sara López, María Sellés, Patricia Rigio, Luis Arechavaleta, Léa Gracia, Roberto Cabria, Yaiza Ontoria, María José Vargas, Jamie McBride, Sofía Capellán Morata, Laura Avivar, Lara Sánchez, Beatriz Rubio, Eva Fonfría y Juan Pablo Izaguirre.
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