Juan Antonio Raga

Juan Antonio Raga


Cavanilles Institute of Biodiversity and Evolutionary Biology,
Science Park, University of Valencia,
Catedratico Josa Beltran 2,
E-46980 Paterna (Valencia), Spain

Department of Zoology, Faculty of Biological Sciences, University of Valencia,
Dr. Moliner 50, E-46100 Burjasot (Valencia), Spain


PhD in Biological Sciences cum laude (with extraordinary award), University of Valencia, Spain.

Professional experience

Current Position

 Professor of Zoology & Director of Science Park (2006-2020) of the University of Valencia, Spain.

Intensive courses

 EU course Marine mammals, seabirds and pollution of marine systems, University of Liege, Belgium; Conferences in many spanish universities. 

Director of the European Seminars on Marine Mammals: Biology and Conservation, Universidad Internacional Menendez Pelayo, Valencia, Spain, 1996-2014; 
Organiser of the 13th Annual Conference of European Cetacean Society, Valencia, Spain, 1999 & the Symposium on Parasites of Marine Mammals in the X International Conference of Parasitology, Vancouver, Canada, 2002; President of the 9th International Symposium on Fish Parasites, University of Valencia-CSIC, Valencia, Spain, 2015.

Awards & Honors

Doctorate extraordinary award of University of Valencia, Spain (1985-1986); University of Valencia-Society Prize awarded to the best Research & Development group of the year (2005); Award to applicate research to Biodiversity Conservation from the “III Premios Biodiversidad y Sociedad” of the Conselleria d'Agricultura, Medi Ambient, Canvi Climàtic i Desenvolupament Rural of the Generalitat Valenciana (2016).


2 scholarships and 7 postdoctoral grants in Spain, France, Denmark, UK, and USA

Editorial Advisor

Diseases of Aquatic Organisms, Editorial Inter-Research (Germany), Foxwell & Davies Scientific Publisher (UK) and Cambridge University Press (UK).

Scientific Editor

Marine Mammals: Biology & Conservation, 2001, Kluwer Academics/Plenum Press, New York, 630 pp, ISBN 0-306-46572-8.

Proyecto mediterraneo. Zonas de especial interes para la conservacion de los cetaceos en el Mediterraneo espanol, 2004, Ministerio de Medio Ambiente. Naturaleza y Parques Nacionales. Serie Tecnica, Madrid, 217 pp, ISBN 84-8014-574-9.

Expert Consultant

  • Reserve Internationale Maritime en Mediterranee Occidentale: Scientific Committee

  • Spanish Ministry of Environment: Marine Species of Wild Flora and Fauna Committee and SAP BIO

  • Spanish Ministry of Agriculture: Fisheries, and Food, Interaction Fisheries-Cetaceans Committee

  • European Commission: Sustainable Marine Ecosystems

  • Agreement on the Conservation of Cetaceans of the Black sea, Mediterranean sea and contiguous Atlantic area: Scientific Committee

  • United Nations Environmental Programme: Regional Activity Center for Specially Protected Areas


Reviewer of Scientific Journals

  • Animal Biodiversity and Conservation (Spain)
  • Aquatic Mammals (USA)
  • Canadian Journal of Fisheries and Aquatic Sciences (Canada)
  • Ciencias Marinas (Mexico)
  • Diseases of Aquatic Organisms (Germany)
  • European Research on Cetaceans (UK)
  • Fishery Bulletin (USA)
  • Folia Parasitologica (Czech Republic)
  • Helminthologia (Slovakia)
  • ICES Journal of Marine Science  (Denmark)
  • Invertebrate Biology (USA)
  • Informes Tecnicos, Spanish Institute of Oceanography (Spain)
  • Journal of the Helminthological Society of Washington (USA)
  • Journal of Parasitology (USA)
  • Journal of Wildlife Diseases (USA)
  • Journal of Zoology (UK)
  • Latin American Journal of Aquatic Mammals (Brazil)
  • Mammalia (France)
  • Mammal Review (UK)
  • Marine Ecology Progress Series (Germany)
  • Marine Mammal Science (USA)
  • NAMMCO Scientific Publications (Norway)
  • Proceedings of the Biological Society of  Washington (USA)
  • Revista Espanola de Herpetologi­a (Spain)
  • Scientia Marina (Spain)
  • The Israeli Journal of Aquaculture (Israel)
  • Scientific Reports of the International Whaling  Commission (UK)
  • Systematic Parasitology (Netherland)
  • Vector-Borne and Zoonotic Diseases (USA)


Referee of Research Applications

  • General Department of Science and Technology & General Department of Hight Education of Ministery of Education and Science, Spain
  • Interministerial Commision of Science and Technology, Spain
  • CODEST committee of European Commission
  • Institut Francais de Recherche pour l'Exploitation de la Mer
  • Fundation Biodiversite de France
  • Agence Nationale de la Recherche, France
  • Agencia Nacional de Promocion Cientifica y Tecnologica (Ministerio de Ciencia, Tecnologi­a e Innovacion Productiva), Republica Argentina
  • Australian Research Council
  • Programa Iberoamericano de Ciencia y Tecnologi­a para el Desarollo (CYTED)
  • Marine Biological Association of the United Kingdom
  • Framework Programmes, European Commission

Research Projects

Participation in 58 competitive research projects (52 as leader) of Spain, France, UK, Italy, Dominican Republic, Tunis, Argentina and EU

Participation in 58 research contracts (as leader) with public and private institutions and enterprises of Spain, Norway and UK


Research Supervisor

Ten Master and 25 Ph D students. Also provided training to a number of foreign students and researchers from Algeria, Argentina, Belgium, Brazil, Belgium, Bulgaria, Chile, Colombia, Ecuador, France, Ireland, Italy, Mexico, Tunis, Uruguay and Ukraine.



 More than 300 scientific papers and book contributions. Some selected recent publications are:

Villar-Torres, M.; Montero, F.; Raga, J.A.; Repullés-Albelda, A.   (2023). The influence of water temperature on the life-cycle of Sparicotyle chrysophrii (Monogenea: Microcotylidae), a common pathogen in gilthead sea bream aquaculture.  Aquaculture, 565, 739103.

Ten, S.; Konishi, K.; Raga, J.A.; Pastene, L.A.; Aznar, F.J. (2022). Epibiotic fauna of the Antarctic minke whale, Balaenoptera bonaerensis, in the Southern Ocean: a reliable indicator of seasonal movements. Scientific Reports, 12, 22214.

Pérez del Olmo, A.; Raga, J.A.; Kostadinova, A. (2022). Parasite communities in a marine fish indicate ecological recovery from the impacts of the Prestige oil-spill 12–13 years after the disaster. Science of the Total Environment, 847: 157354.

Izquierdo-Serrano, M.; Revuelta, O.; Míguez-Lozano, R.; Gozalbes, P.; Ruiz-García, D.; Raga, J.A., Tomás, J. (2022). Assessment of the interactions between cetaceans and fisheries at the south of the Cetacean Migration Corridor and neighboring waters (Western Mediterranean). Frontiers in Marine Science, 1576,

DiMatteo, A.; Cañadas, A.; Roberts, J.; Sparks, L.; Panigada, S.; Boisseau, O.; Moscrop, A.; Fortuna, C.M.; Lauriano, G.; Holcer, D.; Peltier, H.; Ridoux, V.; Raga, J.A.; Tomás, J.; Broderick, A.C.; Godley, B.J.; Haywood, J.; March, D.; Snape, R.; Sagarminaga, R.; Hochscheid, S. (2022). Basin-wide estimates of loggerhead turtle density in the Mediterranean Sea derived from line transect surveys. Frontiers in Marine Science, 1801.

Van Beest, G.S.; Montero, F.E.; Padrós, F.; Raga, J.A.; Born-Torrijos, A. (2022). The versatility of simplicity: structures of Cardiocephaloides longicollis used for different purposes during cercarial transmission. Integrative and Comparative Biology, icac102.

Ten, S.; Raga, J.A.; Aznar, F.J. (2022). Epibiotic fauna on cetaceans worldwide: a systematic review of records and indicator potential. Frontiers in Marine Science, 9:846558.

Pool, R.; Romero-Rubira, C.; Raga, J.A.; Fernández, M.; Aznar, F.J. (2021) Determinants of lungworm specificity in five species of cetacean in the western Mediterranean. Parasites & Vectors, 14: 1-14.

Borrell, A; Gazo, A.; Aguilar, A.; Raga, J.A.; Degollada, E.; Gozalbes, P.; García-Vernet, R. (2021). Niche differentiation amongst northwestern Mediterranean cetaceans using ratios of stable isotopes. Progress in Oceanography, 193: 102559.

Gkafas, G.G.; Jong, M.; Exadactylos, A.; Raga, J.A.; Aznar, F.J.; Hoelzel, A.R. (2020) Sex-specific impact of inbreeding on pathogen load in the striped dolphin. Proceedings of the Royal Society B. -Biological Sciences, 287: 20200195

Cuvertoret-Sanz, M.; López-Figueroa, C.; O’Byrne, A.; Canturri, A.; Martí-Garcia, B.; Pintado, E.; Pérez, L.; Ganges, L.; Cobos, A.; Abarca, M.L.; Raga, J.A.; Van Bressem, M.F.; Domingo, M. (2020). Causes of cetacean stranding and death in the Catalonian coast (Western Mediterranean Sea) in the period 2012-2019.  Diseases of Aquatic Organisms,  142: 239-253 doi:10.3354/dao03550

Novillo, O.; Raga, J.A.; Tomás, J. (2020). Evaluating the presence of microplastics in striped dolphins (Stenella coeruleoalba) stranded in the Western Mediterranean sea. Marine Pollution Bulletin, 160: 111557 doi: 10.1016/j.marpolbul.2020.111557

 Villar-Torres, M.; Repullés-Albelda, A; Montero, F.E.; Raga, J.A.; Blasco Costa, I. (2019). Neither Diplectanum nor specific: A dramatic twist to the taxonomic framework of Diplectanum (Monogenea). International Journal for Parasitology, 49: 365-374

Raga, J.A.; Fernandez, M.; Balbuena, J.A.; Aznar, F.J. (2018). Parasites. In: B. Würsing, J.G.M. Thewissen & K.M. Kovacs (Eds.). Encyclopedia of Marine Mammals, Third Edition, Academic Press/ Elsevier Inc, San Diego, 678-686.

Palacios-Abella, J. F.; Raga, J. A.; Mele, S.; Montero, F. E. (2018). Blood fluke diseases in Pagellus spp. (Sparidae): pathology and description of a new species of Skoulekia in the blackspot seabream P. bogaraveo (Brünnich). Aquaculture, 495: 472–483.

Cañadas, A.; Cotte, C.; Crosti, R.; Fortuna, C.; Gutierrez-Xarxa, R.; Bearzi, G.; Holcer, D.; Raga, J.A.; Rendell, L.; Aissi, M.; Arcangeli, A.; B-Nagy, A.; Campana, I.; Lewis, T.; Laran, S.; Aguilar De Soto, N.; Frantzis, A.; Moulins, A.; Notarbartolo Di Sciara, G.; Pastor, X.; Pulcini, M.; Tepsich, P.; Tomás, J.; Mussi, B.; Politi, E.; Tringali, M.; Azzolin, M.; Di Natale, A.; Rosso, M.; Garcia, P.; Gazo, M.; Lauriano, G.; Panigada, S. (2018). The challenge of habitat modelling for threatened low density species using heterogeneous data: the case of Cuvier's beaked whales in the Mediterranean. Ecological Indicators, 85: 128–136.

Domènech, F.; Tomás, J.; Crespo-Picazo, J.L.; García-Parraga, D.; Raga, J.A.; Aznar F.J.  (2017). To swim or not to swim: potential transmission of Balaenophilus manatorum (Copepoda: Harpacticoida) in marine turtles. PLoS ONE, 12(1): e0170789. doi:10.1371/journal.pone.0170789 

Fraija-Fernández, N.; Fernández, M.; Gozalbes, P.; Revuelta, O.; Raga, J.A.; Aznar F.J. (2017). Living on a harsh habitat: epidemiology of the whale louse, Syncyamus aequus (Cyamidae), in striped dolphins from the western Mediterranean. Journal of Zoology, 303: 199–206, DOI: 10.1111/jzo.12482 

Fraija-Fernández, N.; Fernández, M.; Lehnert, K.; Raga, J.A.; Siebert, U.; Aznar F.J. (2017). Long-distance travellers: phylogeography of a generalist parasite, Pholeter gastrophilus, from Cetaceans. PLoS ONE, 12(1): e0170184. doi:10.1371/journal.pone.0170184  

Palacios-Abella, J.F., Rodríguez-Llanosa, J.; Villora-Montero, M.; Mele, S.; Raga, J.A., Montero, F.E. (2017) Diagnostic accuracy of the light microscope method to detect the eggs of Cardicola spp. in the gill filaments of the bluefin tuna. Veterinary Parasitology, 47: 26-32. 

Hernández-Orts, J.S.; Brandão, M.; Georgieva, S.; Raga, J.A.; Crespo, E.A.; Luque, J.L.; Aznar, F.J. (2017). From mammals back to birds: host-switch of the acanthocephalan Corynosoma australe from pinnipeds to the Magellanic penguin Spheniscus magellanicusPLoS ONE, 12(10): e0183809. 

Gkafas, G.A.; Exadactylos, A; Rogan, E.; Raga, J.A.; Reid, R.; Hoelzel, A.R. (2017). Biogeography and temporal progression during the evolution of striped dolphin population structure in European waters. Journal of Biogeography, 1-11 DOI: 10.1111/jbi.13079. 

Aznar, F.J.; Míguez-Lozano, R.; Ruiz, B., Bosch De Castro, A.; Raga, J.A.; Blanco, C. (2017). Long-term changes (1990-2012) in the diet of striped dolphins Stenella coeruleoalba from the western Mediterranean: a role for overfishing? Marine Ecology Progress Series, 568: 231–247. 

Born-Torrijos, A.; Poulin, R.; Pérez-Del-Olmo, A.; Culurgioni, J.; Raga, J.A.; Holzer, A.S. (2016). An optimized multi-host trematode life cycle: fisheries discards enhance trophic parasite transmission to scavenging birds. International Journal for Parasitology, 46: 745–753 

Clusa, M.; Carreras, C.; Pascual, M.; Gaughran, S.J.; Piovano, S.; Avolio, D.; Ollano, G.; Fernández, G.; Tomás, J.; Raga, J.A., Aguilar, A; Cardona, L. (2016).  Potential bycatch impact on distinct sea turtle populations is dependent on fishing ground rather than gear type in the Mediterranean Sea. Marine Biology, 163:122  doi: 10.1007/s00227-016-2875-1 

Fraija-Fernández, N.; Fernández, M.; Raga, J.A.; Aznar, F.J. (2015). Life-history trade-offs in a generalist digenean from cetaceans: the role of host specificity and environmental factors.  Parasites & Vectors, 8(1): 659

Carrillo, J.M.; Overstreet, R.; Raga, J.A.; Aznar, F.J. (2015). Living on the edge: Settlement patterns by the symbiotic barnacle Xenobalanus globicipitis on small cetaceans. PLoS ONE, 10 (6): e0127367.

Fraija-Fernández, N.; Olson, P.D.; Crespo, E.A.; Raga, J.A.; Aznar, F.; Fernandez, M. (2015). Independent host capture of Cetaceans by Digenean parasites inferred from ribosomal DNA.  International Journal for Parasitology, 45: 167–173. 

Born-Torrijos, A.; Raga, J.A.; Holzer, A. (2015). Trematode maturation patterns in a migratory snail host: What happens during upshore residency in a Mediterranean lagoon? Parasitology Research, 115: 575-585. doi:10.1007/s00436-015-4774-7. 

Revuelta, O.; León, Y.M.; Broderick, A.C..; Feliz, P.; Godley, B.J.; Balbuena, J.A.; Mason, A.; Poulton, K.; Savoré, S.; Raga, J.A.; Tomás, J. (2015). Assessing the efficacy of direct conservation interventions: clutch protection of the leatherback marine turtle in the Dominican Republic. Oryx, 49 (4): 677–686. Doi: 10.1017/S0030605313001488 

Domènech, F.; Badillo, F.J.; Tomás, J.; Raga, J.A.; Aznar, F. (2015). Epibiont communities of loggerhead marine turtles (Caretta caretta) in the western Mediterranean: influence of geographical and ecological factors. Journal of the Marine Biological Association of the United Kingdom, 5(4): 851–861. doi: 10.1017/S0025315414001520. 

Revuelta, O.; Hawkes, L.A.; León, Y.M.; Godley, B.J.; Raga, J.A.; Tomás, J. (2015). Evaluating the importance of Marine Protected Areas for the conservation of hawksbill turtles (Eretmochelys imbricata) nesting in the Dominican Republic. Endangered Species Research, 27: 169–180. 

Ahuir-Baraja, A.E.; Padros, F.; Palacio-Abella, J.F.; Raga, J.A.; Montero, F.E. (2015). Accacoelium contortum (Trematoda: Accacoeliidae) a trematode living as a monogenean: morphological and pathological implications. Parasites & Vectors, 8: 540  doi:10.1186/s13071-015-1162-1 

Mateu, P.;  Nardi, V.; Fraija-Fernández, N.; Mattiucci, S.; Gil De Sola, L.;  Raga, J.A., Fernández, M.; Aznar, F.J (2015). The role of lantern fish (Myctophidae) and cephalopods in the life-cycle of cetacean parasites from western Mediterranean waters. Deep Sea Research I, 95: 115–121. 

Fraija-Fernández, N.; Fernández, M.; Raga, J.A.; Aznar, F.J. (2015). Life-history trade-offs in a generalist digenean from cetaceans: the role of host specificity and environmental factors.  Parasites & Vectors, 8(1): 659  doi: 10.1186/s13071-015-1273-8 

Sánchez-García, N.; Raga, J. A.; Montero, F. E. (2014) Risk assessment for parasites in cultures of Diplodus puntazzo (Sparidae) in the Western Mediterranean: Prospects of cross infection with Sparus aurataVeterinary Parasitology, 204 (3-4): 120-133. 

Revuelta, O. León, Y.M.; Broderick, A.C.; Godley, B.J.; Feliz, P.; Balbuena, J.A.; Raga, J.A.; Tomás, J. (2014) The value of endangered species in protected areas at risk: the case of the leatherback turtle in the Dominican Republic. Biodiversity and Conservation 06/2014; 23(6).

Born-Torrijos, A.; Holzer, A.; Raga, J.A.; Kostadinova, A. (2014) Same host, same lagoon, different transmission pathways: effects of exogenous factors on larval emergence in two marine digenean parasites. Parasitological Research, 113: 545-554.

Mateu, P., Raga, J.A., Fernández, M.; Aznar, F.J. (2014). Intestinal helminth fauna of striped dolphins (Stenella coeruleoalba) in the western Mediterranean: effects of host body length, age and sex. Marine Mammal Science, 30: 961-977.

Born-Torrijos, A.; Poulin, R.; Raga, J.A.; Holzer, A. (2014) Estimating trematode prevalence in snail hosts using a single-step duplex PCR: how badly does cercarial shedding underestimate infection rate? Parasites & Vectors, 7: 243. doi:10.1186/1756-3305-7-243.

Carreras C, Godley BJ, León YM, Hawkes LA, Revuelta O, Raga JA, Tomás J. (2013) Contextualising the Last Survivors: Population Structure of Marine Turtles in the Dominican Republic. PLoS ONE 8(6): e66037. doi:10.1371/journal.pone.0066037.

Hernández-Orts, J.S.; Aznar, F.J.; Blasco-Costa, I.; Garcia, N.A..; Víllora- Montero, M.; Crespo, E.A.; Raga, J.A.; Montero, F.E. (2013). Description, microhabitat selection and infection patterns of sealworm larvae (Pseudoterranova decipiens species complex, Nematoda: Ascaridoidea) in fishes from Patagonia, Argentina. Parasites & Vectors, 6: 1-15 doi:10.1186/1756-3305-6-252.

Revuelta O, León YM, Aznar FJ, Raga JA, Tomás J. (2013) Running against time: conservation of the remaining hawksbill turtle (Eretmochelys imbricata) nesting population in the Dominican Republic. Journal of the Marine Biological Association of the United Kingdom, 93(4): 1133-1140.

Repullés-Albelda, A.; Kostadinova, A.; Raga J. A.; Montero, F. E. (2013) Seasonal population dynamics of Zeuxapta seriolae (Monogenea: Heteraxinidae) parasitizing Seriola dumerili (Carangidae) in the Western Mediterranean. Veterinary Parasitology, 193(1-3): 163-171. 

Quiñones, R.; Giovannini, A.; Raga, J.A.; Fernández, M. (2013). Intestinal helminth fauna of bottlenose dolphin, Tursiops truncatus, and common dolphin, Delphinus delphis, from the western Mediterranean. Journal of Parasitology, 99: 576–579.

Clusa M., Carreras C., Pascual M., Gaughran S.J., Piovano S., Giacoma C., Fernández G., Levy Y., Tomás J., Raga J.A., Maffucci F., Hochscheid S., Aguilar A., Cardona L. (2013). Fine-scale distribution of juvenile Atlantic and Mediterranean loggerhead turtles (Caretta caretta) in the Mediterranean Sea. Marine Biology. DOI 10.1007/s00227-013-2353-y

Alama-Bermejo, G.; Bron, J.E.; Raga J.A.; Holzer, A.S. (2012). 3D morphology, ultrastructure and development of Ceratomyxa puntazzi stages: first insights into the mechanisms of motility and budding in the Myxozoa. PLoS ONE 7(2): e32679. doi:10.1371/journal.pone.0032679.

Sanchez-Garci­a, N., Padros, F., Raga, J.A., Montero, F.E. (2011). Comparative study of the three attachment mechanisms of diplectanid monogeneans. Aquaculture, 318 (3-4): 290-299.

Balbuena, J.A.; Raga, J.A. (2010). Parasites. In: L. Nollet & F. Toldra (Eds.). Handbook of Seafood and Seafood Products Analysis, Chapter 31CRC Press/Taylor & Francis, Boca Raton, 579-602.

Tomas J.; Godley B.J.; Castroviejo J.; Raga J.A (2010). Bioko: Critically  important nesting habitat for sea turtles of West Africa. Biodiversity and Conservation, 19: 2699-2714.

Montero, F.E.; Kostadinova, A.; Raga, J.A. (2009). Development and habitat selection of a new sanguinicolid parasite of cultured greater amberjack, Seriola dumerili, in the Mediterranean. Aquaculture, 288 (1-2): 132-139.

Van Bressem, M-F., Raga, J.A., Di Guardo, G., Jepson, P.D., Duignan, P., Siebert, U., Barrett, T., Santos Mco, Moreno, I.B., Siciliano, S., Aguilar, A. , Van Waerebeek, K. (2009). Emerging infectious diseases in cetaceans worldwide and the possible role of environmental stressors. Diseases of Aquatic Organismsn, 86: 143-157.

Raga, J.A.; Fernandez, M.; Balbuena, J.A.; Aznar, F.J. (2009). Parasites. In: W.F. Perrin, H.G.M. Thewissen & B. Wursing (Eds.). Encyclopedia of Marine Mammals, 2nd Edition, Academic Press, San Diego, 821-830.

Raga, J.A.; Banyard, A.; Domingo, M.; Van Bressem, M.F.; Fernandez, M.; Aznar, F.J.; Barrett, T. (2008). Epizootic of Dolphin Morbillivirus resurges in the Mediterranean. Emerging Infectious Diseases, 14: 471-473.

Gomez de Segura, A.; Hammond, P.S.; Canadas, A.; Raga, J.A. (2007). Comparing cetacean abundance estimates derived from spatial models and design-based line transect methods. Marine Ecology Progress Series, 329: 289-299.

Aznar, F.J.; Fognani, P.; Balbuena, J.A.; Pietrobelli, M.; Raga, J.A. (2006). Distribution patterns of Pholeter gastrophilus (Digenea) in the stomach of four odontocete species: the role of the host digestive physiology. Parasitology, 133: 369-380.