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Universitat De Valencia Facultad De Ciencias Biologicas Tesis UNIVERSITAT DE VALENCIA FACULTAD DE CIENCIAS BIOLOGICAS TESIS DOCTORAL ESTUDIO TAXONOMICO Y ECOLOGICO DE LA PARASITOFAUNA DEL CALDERON COMUN, GLOBICEPHALA HELAS (TRAILL, 1809), EN LAS AGUAS DE EUROPA’ por Juan Antonio Bal buena Diaz-Pinés Di rector Juan Antonio Raga Esteve Valencia junio, 1991 * Trabajo encuadrado en el proyecto PB87-997 de la DGICYT UMI Number: U607653 All rights reserved INFORMATION TO ALL USERS The quality of this reproduction is dependent upon the quality of the copy submitted. In the unlikely event that the author did not send a complete manuscript and there are missing pages, these will be noted. Also, if material had to be removed, a note will indicate the deletion. Disscrrlation Publishíng UMI U607653 Published by ProQuest LLC 2014. Copyright in the Dissertation held by the Author. Microform Edition © ProQuest LLC. All rights reserved. This work is protected against unauthorized copying underTitle 17, United States Code. ProQuest LLC 789 East Eisenhower Parkway P.O. Box 1346 Ann Arbor, MI 48106-1346 Dr. Moliner, 50 46100 - BURJASOT (Valencia) España UNIVERSITAT DE VALENCIA DEPARTAMENT DE BIOLOGIA ANIMAL, BIOLOGIA CEL.LULAR. I PARASITOLOGIA D- Juan Antonio Raga Esteve, Profesor Titular de Biología Animal de la Universitat de Valéncia, CERTIFICA que el trabajo titulado “Estudio Taxonómico y Ecológico de la Parasitofaúna del Calderón Común, Clobicephala melas (Traill, 1809), en las Aguas de Europa'*, que presenta D. Juan Antonio Ealbuena Díaz—Pinés para optar al grado de Doctor en Ciencias Biológicas, ha sido realizado en el Departamento de Biología Animal, Biología Celular y Parasitología de la Universitat de Valéncia bajo mi dirección. Burjassot, E7 de mayo de 1991 ¿fe - Juan Antonio Esteve Pf.JWOSW R ?í/3 Taxonomic and Ecológica! Study of the Parasite Fauna of the Long- finned Pilot Whale, Globicephala me Tas (Traill, 1809), in European Waters. ABSTRACT.- A study of the taxonomy and ecology of parasi tes and epizoics of long-finned pilot whales (Globicephala melas) from three European regions (Faroe Islands, French Atlantic and Spanish Mediterranean) has been carried out. In all, 21 (four crustacean and 21 helminth) species were detected. Two of them, Hadwenius delamurei and Crassicauda carbonel1 i, are new to Science. The finding of Hadwenius subtilus, Orthosplanchus albamarinus, Diphyllobothrium polyrugosum and D. stemmacephalum in the long- finned pilot whale constitute new host records. An investigation focussing on the egg size of C. carbonel Ti, evidenced that egg length and width have little valué in the taxonomy of the genus Crassicauda, as previously suggested by Dollfus (1968a). A comparison of the helminth faunal compositions showed differences across the three areas studied. Differences between the helminth fauna of the Faroese pilot whales and that from pilot whales in Newfoundland waters were al so observed. These geographical variations are possibly due to several factors, such as sampling methodology, environmental conditions (specially water temperature) and intermediate-host fauna in the regions concerned. The prevalence, abundance and spatial distribution of the whale lice Isocyamus delphini on Faroese pilot whales showed significant differences between adult males and the rest of individuáis. Behavioural factors seemed to account for these results. Host age was the most important factor determining the prevalence of the stomach helminth species (Anisakis simplex and Pholeter gastrophilus) from pilot whales off the Faroes, animals > 3 yrs oíd showing higher prevalences. First infections were detected as soon as unweaned whales started their preying activities. A. simplex was the commonest of all species detected at the Faroe Islands. This is possibly due to its low specificity for both the intermediate and definitive hosts. In addition, previous studies showed a high abundance of this species in different intermediate hosts in the North Atlantic. The intestinal helminth infracommunities of the pilot whales taken at the Faroes appeared to be depaupérate and ai solacionist in character. However, some features indicated potential interactions between the congeners H. delamurei and H. subtilus. The abundance of the component intestinal species (prevalence > 10 %) of the Faroese pilot whales was significantly higher in animals >. 10 yrs oíd than in younger individuáis, the former showing a more, diverse helminth community as well. A higher food intake among whales >_ 10 yrs oíd related to body size may account for these results. Differences in helminth abundances between host sexes may be due to (i) physiologycal factors and (ii) differences in body size between whale genders. Likewise, the abundance of the helminth species varied across schools. Our results are consistent with previous studies of organochlorine contents in blubber, supporting the idea of several sympatric pilot whale populations in Faroese waters. These differences might al so be due to some extent to seasonal or long-term variations of the abundance of infective stages. Eco-evolutive factors seemeed to be more impotant than host phylogeny in determining the structure of the intestinal helminth comunity of the long-finned pilot whale. i i INDICE AGRADECIMIENTOS....................................................... v 1 .-INTRODUCCION....................................................... 1 1.1. El Parasitismo y su Importancia como Asociación Biológica.............................. 2 1.2. El Parasitismo en el Medio Marino.................... 7 1.3. Biología del Calderón Común.......................... 10 1.4. Justificación y Objetivos del Trabajo..............12 2.- MATERIAL Y METODOS............................................. 14 2.1. Obtención de los Hospedadores........................15 2.2. Necropsias y Recolección del Material..............21 2.3. Métodos Parasitológicos............................... 25 2.4. Obtención de Datos Biológicos de los Hospedadores.....................................30 2.5. Métodos Analíticos y Terminología empleada........31 3.- RESULTADOS Y DISCUSION........................................ 43 3.1. Estudio Taxonómico.......... 44 3.1.1. Cirripedia.......................................45 Conchoderma auritum............................46 Xenobalanus globicipitis......................54 3.1.2. Malacostraca.....................................60 Isocyamus delphini............................. 61 3.1.3. Di genea...........................................69 Hadwenius delamurei............................70 Hadweni us subtilus............................. 85 Orthosplanchnus albamariñus.................. 97 Pholeter gastrophi lus........................ 104 3.1.4. Cestoda............ 110 Monorygma gr i mal di i...........................111 Phyl lobothrium delphini......................118 Trigonocotyle globicephalae.................1 23 i i i Diphyllobothrium s p . A ....................... 129 Diphyllobothrium sp. B ....................... 142 Diphyl lobothrium polyrugosum.............. .152 Diphyl lobothr i um stemmacephal um. .......... 160 Diphyllobothriidae sp ........................ 168 3.1.5. Acantocephala..................................173 Bolbosoma capitatum...........................174 3.1.6. Nematoda........................................180 Anisakis simplex...............................181 Crassicauda carbonelli....................... 189 Stenurus globicephalae....................... 201 3.2. Estudio Ecológico...................................... 210 3.2.1. Parasitofauna en las Feroe - Comparación con Otras Areas.................211 3.2.2. Isocyamus delphini: Relaciones Parási to-Hospedador...........................218 Resul tados...................................... 218 Discusión ................................219 3.2.3. Helmintos Estomacales........................226 3.2.4. Helmintos Intestinales...................... 229 Resul tados...................................... 229 Di scusi ón ....................................... 252 3.3. Patologías Parasitarias...............................278 4 . -RESUMEN Y CONCLUSIONES................................. 282 BIBIOGRAFIA..........................................................288 i v AGRADECIMIENTOS En primer lugar, debo expresar mi más profunda gratitud al Dr. Juan Antonio Raga Esteve, mi director durante estos últimos años, por su paciencia, dedicación y valiosos consejos en la preparación de este estudio. El trabajo en las islas Feroe fue posible gracias a la Sra. Dórete Bloch del Foroya Náttúrugripasvn. A ella, le agradezco las innumerables molestias que se tomó realizando distintas gestiones que facilitaron grandemente este trabajo. Igualmente, le doy las gracias por su interés e inestimable ayuda durante el trabajo de campo. Asimismo, estoy profundamente agradecido a la Dra. Geneviéve Desportes, de la citada institución, por sus consejos, por su valiosa asistencia en el muestreo y por su amistad y apoyo. Ambas me facilitaron, además, una gran cantidad de datos que posibilitaron gran parte de este trabajo. Al Dr. Jóhannes Jóhansen, Director del F0roya Náttúrugripasvn, doy las gracias por su hospitalidad al dejarme utilizar las instalaciones del museo que dirige, por su infinita paciencia y por su interés en este estudio. Igualmente, doy mis más expresivas gracias al personal del museo por su asistencia; muy especialmente, al Sr. Rógvi Mouritsen, por su valiosísima ayuda técnica y por honrarme con su amistad. También debo destacar la labor de S. Skaaning, de E. Stefansson y de H. Gaasedal. El trabajo de campo
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