{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,17]],"date-time":"2026-07-17T21:16:51Z","timestamp":1784323011068,"version":"3.55.0"},"reference-count":122,"publisher":"MDPI AG","issue":"14","license":[{"start":{"date-parts":[[2025,7,11]],"date-time":"2025-07-11T00:00:00Z","timestamp":1752192000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"LIFE Program of the European Union","award":["LIFE PINNA LIFE20 NAT\/IT\/001122 CUP B95F21001880002"],"award-info":[{"award-number":["LIFE PINNA LIFE20 NAT\/IT\/001122 CUP B95F21001880002"]}]},{"name":"LIFE Program of the European Union","award":["CN00000033"],"award-info":[{"award-number":["CN00000033"]}]},{"name":"Italian Ministry of University and Research, PNRR","award":["LIFE PINNA LIFE20 NAT\/IT\/001122 CUP B95F21001880002"],"award-info":[{"award-number":["LIFE PINNA LIFE20 NAT\/IT\/001122 CUP B95F21001880002"]}]},{"name":"Italian Ministry of University and Research, PNRR","award":["CN00000033"],"award-info":[{"award-number":["CN00000033"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Animals"],"abstract":"<jats:p>This review addresses the critical conservation challenges faced by Pinna nobilis, the noble pen shell, a keystone umbrella species in Mediterranean marine ecosystems. Since 2016, the species has experienced catastrophic population declines due to mass mortality events likely driven by protozoan, bacterial, and viral infections. Despite these severe circumstances, small resilient populations persist in select estuaries and coastal lagoons across the Mediterranean, offering potential for recovery. We provide a comprehensive overview on research dedicated to Pinna nobilis\u2019 biology, genetic variation, disease dynamics, and environmental factors influencing its survival, with a focus on refugia where populations still endure. Remarkably, recent studies have revealed signs of resistance in certain individuals and the potential for hybridisation with Pinna rudis. In this context, the possible impact of the increasing occurrence of hybridisation between Pinna nobilis and Pinna rudis on the conservation of their genetic diversity should be carefully considered. This review highlights the importance of ongoing conservation efforts including habitat restoration, protection of remaining populations, assessment of past and present genetic variability, and the development of captive breeding programmes. We aim to elucidate the need for continued studies on Pinna nobilis\u2019 biodiversity, particularly its evolutionary dynamics, genetic makeup, and the interplay of environmental variables influencing its survival and persistence.<\/jats:p>","DOI":"10.3390\/ani15142044","type":"journal-article","created":{"date-parts":[[2025,7,11]],"date-time":"2025-07-11T10:26:53Z","timestamp":1752229613000},"page":"2044","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":7,"title":["Pinna nobilis, the Vanishing Giant: A Comprehensive Review on the Decline of a Mediterranean Icon"],"prefix":"10.3390","volume":"15","author":[{"given":"Ilenia","family":"Azzena","sequence":"first","affiliation":[{"name":"Department of Veterinary Medicine, University of Sassari, Via Vienna 2, 07100 Sassari, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chiara","family":"Locci","sequence":"additional","affiliation":[{"name":"Department of Veterinary Medicine, University of Sassari, Via Vienna 2, 07100 Sassari, Italy"},{"name":"Department of Biomedical Sciences, University of Sassari, Viale San Pietro 43b, 07100 Sassari, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Noemi","family":"Pascale","sequence":"additional","affiliation":[{"name":"Department of Veterinary Medicine, University of Sassari, Via Vienna 2, 07100 Sassari, Italy"},{"name":"Department of Biomedical Sciences, University of Sassari, Viale San Pietro 43b, 07100 Sassari, Italy"},{"name":"Department of Chemical, Physical, Mathematical, and Natural Sciences, University of Sassari, Via Vienna 2, 07100 Sassari, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0008-3657-8740","authenticated-orcid":false,"given":"Ilaria","family":"Deplano","sequence":"additional","affiliation":[{"name":"Department of Biomedical Sciences, University of Sassari, Viale San Pietro 43b, 07100 Sassari, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Riccardo","family":"Senigaglia","sequence":"additional","affiliation":[{"name":"Department of Veterinary Medicine, University of Sassari, Via Vienna 2, 07100 Sassari, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3501-714X","authenticated-orcid":false,"given":"Fabio","family":"Scarpa","sequence":"additional","affiliation":[{"name":"Department of Biomedical Sciences, University of Sassari, Viale San Pietro 43b, 07100 Sassari, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5042-4851","authenticated-orcid":false,"given":"Marco","family":"Casu","sequence":"additional","affiliation":[{"name":"Department of Veterinary Medicine, University of Sassari, Via Vienna 2, 07100 Sassari, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3058-4663","authenticated-orcid":false,"given":"Daria","family":"Sanna","sequence":"additional","affiliation":[{"name":"Department of Biomedical Sciences, University of Sassari, Viale San Pietro 43b, 07100 Sassari, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2025,7,11]]},"reference":[{"key":"ref_1","unstructured":"Garcia-March, J.R. (2005). Aportaciones al Conocimiento de la Biolog\u00eda de Pinna nobilis Linneo, 1758 (Mollusca: Bivalvia) en el Litoral Mediterr\u00e1neo Ib\u00e9rico. [Ph.D. Thesis, Universitat de Valencia]."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"667","DOI":"10.12681\/mms.836","article-title":"Utilization of muddy detritus as organic matter source by the fan mussel Pinna nobilis","volume":"15","author":"Trigos","year":"2014","journal-title":"Mediterr. Mar. Sci."},{"key":"ref_3","first-page":"1","article-title":"From dark to light and back again: Is Pinna nobilis, the largest Mediterranean shellfish, on the brink of extinction? What about Pinna nobilis","volume":"70","author":"Scarpa","year":"2021","journal-title":"Veterinaria"},{"key":"ref_4","doi-asserted-by":"crossref","unstructured":"V\u00e1zquez-Luis, M., \u00c1lvarez, E., Barraj\u00f3n, A., Garc\u00eda-March, J.R., Grau, A., Hendriks, I.E., Jim\u00e9nez, S., Kersting, D., Moreno, D., and P\u00e9rez, M. (2017). SOS Pinna nobilis: A mass mortality event in western Mediterranean Sea. Front. Mar. Sci., 4.","DOI":"10.3389\/fmars.2017.00220"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1016\/j.jip.2018.07.006","article-title":"Haplosporidium pinnae sp. nov.; a haplosporidan parasite associated with mass mortalities of the fan mussel, Pinna nobilis, in the Western Mediterranean Sea","volume":"157","author":"Catanese","year":"2018","journal-title":"J. Invertebr. Pathol."},{"key":"ref_6","doi-asserted-by":"crossref","unstructured":"Carella, F., Aceto, S., Pollaro, F., Miccio, A., Iaria, C., Carrasco, N., Prado, P., and De Vico, G. (2019). A mycobacterial disease is associated with the silent mass mortality of the pen shell Pinna nobilis along the Tyrrhenian coastline of Italy. Sci. Rep., 9.","DOI":"10.1038\/s41598-018-37217-y"},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Carella, F., Prado, P., De Vico, G., Pali\u0107, D., Villari, G., Garc\u00eda-March, J.R., Tena-Medialdea, J., Cort\u00e9s-Melendreras, E., Gim\u00e9nez-Casalduero, F., and Sigovini, M. (2023). A widespread picornavirus affects the hemocytes of the noble pen shell (Pinna nobilis), leading to its immunosuppression. Front. Vet. Sci., 10.","DOI":"10.3389\/fvets.2023.1273521"},{"key":"ref_8","unstructured":"Kersting, D., Benabdi, M., \u010ci\u017emek, H., Grau, A., Jimenez, C., Katsanevakis, S., \u00d6zt\u00fcrk, B., Tuncer, S., Tunesi, L., and V\u00e1zquez-Luis, M. (2019). Pinna nobilis. The IUCN Red List of Threatened Species 2019\/\/IUCN Red List of Threatened Species. IUCN, e.T160075998A160081499."},{"key":"ref_9","first-page":"164","article-title":"Revision of the worldwide Recent Pinnidae and some remarks on fossil European Pinnidae","volume":"13","author":"Schultz","year":"2013","journal-title":"Acta Conchyl."},{"key":"ref_10","unstructured":"Huber, M. (2010). Compendium of Bivalves. A Full-Color Guide to 3\u2019300 of the World\u2019s Marine Bivalves. A Status on Bivalvia After 250 Years of Research, ConchBooks."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1016\/j.ympev.2014.02.008","article-title":"First molecular phylogeny of the circumtropical bivalve family Pinnidae (Mollusca, Bivalvia): Evidence for high levels of cryptic species diversity","volume":"75","author":"Lemer","year":"2014","journal-title":"Mol. Phylogenet. Evol."},{"key":"ref_12","first-page":"37","article-title":"Ecology of the pterioid bivalves Pinna bicolor Gmelin and Pinna nobilis L.","volume":"3","author":"Butler","year":"1993","journal-title":"Mar. Life"},{"key":"ref_13","first-page":"93","article-title":"Contribution to the ecology of Pinna nobilis L. (Moll.; Bivalvia) in the northern Adriatic","volume":"3","author":"Zavodnik","year":"1967","journal-title":"Thalass. Jugosl."},{"key":"ref_14","unstructured":"Boudouresque, C.F., Avon, M., and Gravez, V. (1991). Les Esp\u00e8ces Marines \u00e0 Prot\u00e9ger en M\u00e9diterran\u00e9e, GIS Posidonie publications."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"51","DOI":"10.3354\/esr001051","article-title":"Population ecology of the endangered fan mussel Pinna nobilis in a marine lake","volume":"1","author":"Katsanevakis","year":"2005","journal-title":"Endanger. Species Res."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"109","DOI":"10.1016\/bs.amb.2015.06.002","article-title":"The pen shell, Pinna nobilis: A review of population status and recommended research priorities in the Mediterranean Sea","volume":"71","author":"Basso","year":"2015","journal-title":"Adv. Mar. Biol."},{"key":"ref_17","first-page":"2779","article-title":"The byssus threads of Pinna nobilis: A histochemical and ultrastructural study","volume":"61","author":"Diana","year":"2017","journal-title":"Eur. J. Histochem."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"335","DOI":"10.1098\/rstb.1953.0006","article-title":"Form and habit in Pinna carnea Gmelin","volume":"237","author":"Yonge","year":"1953","journal-title":"Philos. Trans. R. Soc. Lond. Soc. B"},{"key":"ref_19","first-page":"175","article-title":"The family PINNIDAE in the Indo-Pacific","volume":"1","author":"Rosewater","year":"1961","journal-title":"Indo-Pac. Mollusca"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"621","DOI":"10.1666\/06-096.1","article-title":"The duplivincular ligament of recent Pinna nobilis L.; 1758: Further evidence for pterineid ancestry of the Pinnoidea","volume":"82","author":"Carter","year":"2008","journal-title":"J. Paleontol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"689","DOI":"10.1007\/s00227-007-0842-6","article-title":"Shell gaping behaviour of Pinna nobilis L.; 1758: Circadian and circalunar rhythms revealed by in situ monitoring","volume":"153","year":"2008","journal-title":"Mar. Biol."},{"key":"ref_22","first-page":"209","article-title":"From youth to death of old age: The 50-year story of a Pinna nobilis fan mussel population at Port-Cros Island (Port-Cros National Park, Provence, Mediterranean Sea)","volume":"29","author":"Rouanet","year":"2015","journal-title":"Sci. Rep. Port-Cros Natl. Park"},{"key":"ref_23","unstructured":"Czihak, G., and Dierl, W. (1961). Pinna nobilis L.: Eine Pr\u00e4parationsanleitung, Gustav Fisher."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"246","DOI":"10.1016\/j.ecss.2010.12.033","article-title":"Size-differential feeding in Pinna nobilis L. (Mollusca: Bivalvia): Exploitation of detritus, phytoplankton and zooplankton. Estuar","volume":"92","author":"Davenport","year":"2011","journal-title":"Coast. Shelf Sci."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"77","DOI":"10.1016\/j.jembe.2015.03.006","article-title":"Evaluating stable isotopic signals in bivalve Pinna nobilis under different human pressures","volume":"467","author":"Alomar","year":"2015","journal-title":"J. Exp. Mar. Biol. Ecol."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"921","DOI":"10.1007\/s00227-012-2144-x","article-title":"Diets of fan shells (Pinna nobilis) of different sizes: Fatty acid profiling of digestive gland and adductor muscle","volume":"160","author":"Najdek","year":"2013","journal-title":"Mar. Biol."},{"key":"ref_27","first-page":"99","article-title":"Successive hermaphrodism with asynchronous maturation of Pinna nobilis (L.) (Bivalvia: Pterioidea)","volume":"318","year":"1995","journal-title":"CR Acad. Sci. III"},{"key":"ref_28","unstructured":"Giese, A.C., and Pearse, J.S. (1979). Reproduction of Marine Invertebrates, Academic Press Inc."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"205","DOI":"10.1007\/s002270050459","article-title":"Age and growth of the fan mussel Pinna nobilis from south-east Spanish Mediterranean seagrass (Posidonia oceanica) meadows","volume":"133","author":"Richardson","year":"1999","journal-title":"Mar. Biol."},{"key":"ref_30","first-page":"83","article-title":"Ecologie de Pinna nobilis (l.) mollusque bivalve sur les c\u00f4tes de corse. Essais de transplantation et exp\u00e9riences en milieu contr\u00f4l\u00e9","volume":"20","author":"Vicente","year":"1990","journal-title":"Haliotis"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"282","DOI":"10.1016\/j.marenvres.2017.08.007","article-title":"Long-term assessment of recruitment, early stages and population dynamics of the endangered Mediterranean fan mussel Pinna nobilis in the Columbretes Islands (NW Mediterranean)","volume":"130","author":"Kersting","year":"2017","journal-title":"Mar. Environ. Res."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"823","DOI":"10.1017\/S0025315410001694","article-title":"On the functional morphology of Pinna and Atrina larvae (Bivalvia: Pinnidae) from the Atlantic","volume":"91","author":"Allen","year":"2011","journal-title":"J. Mar. Biol. Assoc. United Kingd."},{"key":"ref_33","first-page":"115","article-title":"Early life-history of useful bivalves in the Ariake Sea (1)","volume":"6","author":"Yoshida","year":"1956","journal-title":"J. Shimonoseki Univ. Fish."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"107","DOI":"10.2331\/suisan.27.107","article-title":"Identification of the larva of Pinna atrina japonica (Reeve)","volume":"27","author":"Ota","year":"1961","journal-title":"Bull. Jpn. Soc. Sci. Fish."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"2171","DOI":"10.4319\/lo.2003.48.6.2171","article-title":"Climate-related changes in recruitment of the bivalve Macoma balthica","volume":"48","author":"Philippart","year":"2003","journal-title":"Limnol. Oceanogr."},{"key":"ref_36","first-page":"25","article-title":"Modelling the recruitment effect in a small marine protected area: The example of saltwater lakes on the Island of Mljet (Adriatic Sea)","volume":"49","author":"Peharda","year":"2008","journal-title":"Acta Adriat."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"1385","DOI":"10.1080\/00222930110056892","article-title":"Epibiontic mollusc communities on Pinna nobilis L. (Bivalvia, Mollusca)","volume":"36","author":"Giacobbe","year":"2002","journal-title":"J. Nat. Hist."},{"key":"ref_38","unstructured":"Cosentino, A., and Giacobbe, S. (2008). Aspects of epizoobiontic mollusc assemblages on Pinna shells. II. Does the Mediterranean P. nobilis represent an isle of biodiversity?. Cah. Biol. Mar., 49."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"129","DOI":"10.3989\/scimar.2009.73n1129","article-title":"Associated fauna of the fan shell Pinna nobilis (Mollusca: Bivalvia) in the northern and eastern Tunisian coasts","volume":"73","author":"Rabaoui","year":"2009","journal-title":"Sci. Mar."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"143","DOI":"10.3989\/scimar.2009.73n1143","article-title":"Density, size structure, shell orientation and epibiontic colonization of the fan mussel Pinna nobilis L. 1758 (Mollusca: Bivalvia) in three contrasting habitats in an estuarine area of Sardinia (W Mediterranean)","volume":"73","author":"Addis","year":"2009","journal-title":"Sci. Mar."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"1227","DOI":"10.1017\/S0025315400038741","article-title":"The occurrence of Pontonia pinnophylax (Decapoda: Natantia: Pontoninae) in Pinna nobilis (Mollusca: Bivalvia: Pinnidae) from the Mediterranean","volume":"77","author":"Richardson","year":"1997","journal-title":"J. Mar. Biol. Assoc. United Kingd."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"433","DOI":"10.1163\/156854008783797507","article-title":"Two species of Crustacea (Decapoda) associated with the fan mussel, Pinna nobilis Linnaeus, 1758 (Mollusca, Bivalvia)","volume":"81","author":"Rabaoui","year":"2008","journal-title":"Crustaceana"},{"key":"ref_43","doi-asserted-by":"crossref","unstructured":"Scarpa, F., Sanna, D., Azzena, I., Mugetti, D., Cerruti, F., Hosseini, S., Cossu, P., Pinna, S., Grech, D., and Cabana, D. (2020). Multiple non-species-specific pathogens possibly triggered the mass mortality in Pinna nobilis. Life, 10.","DOI":"10.3390\/life10100238"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"339","DOI":"10.1007\/s10661-006-9479-z","article-title":"Monitoring of the endangered Pinna nobilis Linne, 1758 in the Mar Grande of Taranto (Ionian sea, Italy)","volume":"131","author":"Centoducati","year":"2007","journal-title":"Environ. Monit. Assess."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"340","DOI":"10.1093\/mollus\/eyq023","article-title":"Modelling population density of Pinna nobilis (Bivalvia) on the eastern and southeastern coast of Tunisia","volume":"76","author":"Rabaoui","year":"2010","journal-title":"J. Molluscan Stud."},{"key":"ref_46","unstructured":"Raphael, M. (2010). The Cruising Chef Cookbook, Paradise Cay Publications."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"786","DOI":"10.1071\/MF14285","article-title":"Influence of boat anchoring on Pinna nobilis: A field experiment using mimic units","volume":"66","author":"Borg","year":"2015","journal-title":"Mar. Freshw. Res."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"31","DOI":"10.1002\/aqc.3237","article-title":"Human impacts on the endangered fan mussel, Pinna nobilis","volume":"30","author":"Kaiser","year":"2020","journal-title":"Aquat. Conserv. Mar. Freshw. Ecosyst."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"e12607","DOI":"10.1111\/maec.12607","article-title":"Mass mortality of the fan mussel, Pinna nobilis in Turkey (eastern Mediterranean)","volume":"41","author":"Alan","year":"2020","journal-title":"Mar. Ecol."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"429","DOI":"10.12681\/mms.42","article-title":"\u201cProtected\u201d marine shelled molluscs: Thriving in Greek seafood restaurants","volume":"12","author":"Katsanevakis","year":"2011","journal-title":"Mediterr. Mar. Sci."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"560","DOI":"10.1177\/03009858231186737","article-title":"Multipathogen infections and multifactorial pathogenesis involved in noble pen shell (Pinna nobilis) mass mortality events: Background and current pathologic approaches","volume":"60","author":"Carella","year":"2023","journal-title":"Vet. Pathol."},{"key":"ref_52","doi-asserted-by":"crossref","unstructured":"Grau, A., Villalba, A., Navas, J.I., Hansjosten, B., Valencia, J.M., Garc\u00eda-March, J.R., Prado, P., Follana-Bern\u00e1, G., Morage, T., and V\u00e1zquez-Luis, M. (2022). Wide-geographic and long-term analysis of the role of pathogens in the decline of Pinna nobilis to critically endangered species. Front. Mar. Sci., 9.","DOI":"10.3389\/fmars.2022.666640"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"14","DOI":"10.1016\/j.jip.2017.05.006","article-title":"First haplosporidan parasite reported infecting a member of the Superfamily Pinnoidea (Pinna nobilis) during a mortality event in Alicante (Spain, Western Mediterranean)","volume":"148","author":"Darriba","year":"2017","journal-title":"J. Invertebr. Pathol."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"150","DOI":"10.3391\/ai.2019.14.2.01","article-title":"The cryptogenic parasite Haplosporidium pinnae invades the Aegean Sea and causes the collapse of Pinna nobilis populations","volume":"14","author":"Katsanevakis","year":"2019","journal-title":"Aquat. Invasions"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"32","DOI":"10.1016\/j.jip.2019.04.005","article-title":"Haplosporidium pinnae associated with mass mortality in endangered Pinna nobilis (Linnaeus 1758) fan mussels","volume":"164","author":"Panarese","year":"2019","journal-title":"J. Invertebr. Pathol."},{"key":"ref_56","doi-asserted-by":"crossref","unstructured":"Cabanellas-Reboredo, M., V\u00e1zquez-Luis, M., Mourre, B., \u00c1lvarez, E., Deudero, S., Amores, \u00c1., Addis, P., Ballesteros, E., Barraj\u00f3n, A., and Coppa, S. (2019). Tracking a mass mortality outbreak of pen shell Pinna nobilis populations: A collaborative effort of scientists and citizens. Sci. Rep., 9.","DOI":"10.1038\/s41598-019-49808-4"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"104889","DOI":"10.1016\/j.marenvres.2020.104889","article-title":"First detection of the invasive Haplosporidian and Mycobacteria parasites hosting the endangered bivalve Pinna nobilis in Thermaikos Gulf, North Greece","volume":"155","author":"Lattos","year":"2020","journal-title":"Mar. Environ. Res."},{"key":"ref_58","doi-asserted-by":"crossref","unstructured":"\u0160ari\u0107, T., \u017dupan, I., Aceto, S., Villari, G., Pali\u0107, D., De Vico, G., and Carella, F. (2020). Epidemiology of noble pen shell (Pinna nobilis L. 1758) mass mortality events in Adriatic Sea is characterised with rapid spreading and acute disease progression. Pathogens, 9.","DOI":"10.3390\/pathogens9100776"},{"key":"ref_59","first-page":"221","article-title":"Is Pinna nobilis L. critically endangered in Neum bay, Bosnia and Herzegovina?","volume":"69","year":"2020","journal-title":"Veterinaria"},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"450","DOI":"10.1080\/1354750X.2021.1910344","article-title":"Mass mortality in endangered fan mussels Pinna nobilis (Linnaeus 1758) caused by co-infection of Haplosporidium pinnae and multiple Vibrio infection in \u00c7anakkale Strait, Turkey","volume":"26","author":"Aksu","year":"2021","journal-title":"Biomarkers"},{"key":"ref_61","doi-asserted-by":"crossref","unstructured":"L\u00f3pez-Sanmart\u00edn, M., Catanese, G., Grau, A., Valencia, J.M., Garc\u00eda-March, J.R., and Navas, J.I. (2019). Real-Time PCR based test for the early diagnosis of Haplosporidium pinnae affecting fan mussel Pinna nobilis. PLoS ONE, 14.","DOI":"10.1371\/journal.pone.0212028"},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"107735","DOI":"10.1016\/j.jip.2022.107735","article-title":"New insights about Haplosporidium pinnae and the pen shell Pinna nobilis mass mortality events","volume":"190","author":"Tiscar","year":"2022","journal-title":"J. Invertebr. Pathol."},{"key":"ref_63","doi-asserted-by":"crossref","unstructured":"Carella, F., Antuofermo, E., Farina, S., Salati, F., Mandas, D., Prado, P., Panarese, R., Marino, F., Fiocchi, E., and Pretto, T. (2020). In the wake of the ongoing mass mortality events: Co-occurrence of Mycobacterium, Haplosporidium and other pathogens in Pinna nobilis collected in Italy and Spain (Mediterranean Sea). Front. Mar. Sci., 7.","DOI":"10.3389\/fmars.2020.00382"},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"734899","DOI":"10.1016\/j.aquaculture.2019.734899","article-title":"Presence of Vibrio mediterranei associated to major mortality in stabled individuals of Pinna nobilis L.","volume":"519","author":"Prado","year":"2020","journal-title":"Aquaculture"},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"109664","DOI":"10.1016\/j.fsi.2024.109664","article-title":"Measuring immunocompetence in the natural population and captive individuals of noble pen shell Pinna nobilis affected by Pinna nobilis Picornavirus (PnPV)","volume":"151","author":"Carella","year":"2024","journal-title":"Fish Shellfish Immunol."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"581","DOI":"10.1007\/s10661-020-08541-6","article-title":"Assessment of predominant bacteria in noble pen shell (Pinna nobilis) collected in the Eastern Adriatic Sea","volume":"192","author":"Pavlinec","year":"2020","journal-title":"Environ. Monit. Assess."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"109202","DOI":"10.1016\/j.ecss.2025.109202","article-title":"The mass mortality of Pinna nobilis throughout the Mediterranean Sea has not yet affected the genetic diversity of the species in one of the last genetic reservoirs","volume":"317","author":"Peyran","year":"2025","journal-title":"Estuar. Coast. Shelf Sci."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"15261","DOI":"10.1073\/pnas.0403809101","article-title":"Most species are not driven to extinction before genetic factors impact them","volume":"101","author":"Spielman","year":"2004","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"704","DOI":"10.1038\/s41586-024-08458-x","article-title":"Global meta-analysis shows action is needed to halt genetic diversity loss","volume":"638","author":"Shaw","year":"2025","journal-title":"Nature"},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"1293","DOI":"10.1007\/s10709-012-9631-3","article-title":"Patterns of spatial genetic structuring in the endangered limpet Patella ferruginea: Implications for the conservation of a Mediterranean endemic","volume":"139","author":"Casu","year":"2011","journal-title":"Genetica"},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"412","DOI":"10.2478\/s11756-008-0061-8","article-title":"Genetic structure of the endangered species Pinna nobilis (Mollusca: Bivalvia) inferred from mtDNA sequences","volume":"63","author":"Katsares","year":"2008","journal-title":"Biologia"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"99","DOI":"10.1007\/s10750-011-0827-9","article-title":"Genetic variation among populations of the endangered fan mussel Pinna nobilis (Mollusca: Bivalvia) along the Tunisian coastline","volume":"678","author":"Rabaoui","year":"2011","journal-title":"Hydrobiologia"},{"key":"ref_73","doi-asserted-by":"crossref","unstructured":"Sanna, D., Cossu, P., Dedola, G.L., Scarpa, F., Maltagliati, F., Castelli, A., Franzoi, P., Lai, T., Cristo, B., and Curini-Galletti, M. (2013). Mitochondrial DNA reveals genetic structuring of Pinna nobilis across the Mediterranean Sea. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0067372"},{"key":"ref_74","first-page":"31","article-title":"Highly polymorphic microsatellite markers for the Mediterranean endemic fan mussel Pinna nobilis","volume":"16","author":"Costa","year":"2015","journal-title":"Mediterr. Mar. Sci."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"175","DOI":"10.1002\/aqc.2976","article-title":"Gene pool and connectivity patterns of Pinna nobilis in the Balearic Islands (Spain, Western Mediterranean Sea): Implications for its conservation through restocking","volume":"29","author":"Basso","year":"2019","journal-title":"Aquat. Conserv. Mar. Freshw. Ecosyst."},{"key":"ref_76","doi-asserted-by":"crossref","unstructured":"Wesselmann, M., Gonz\u00e1lez-Wang\u00fcemert, M., Serr\u00e3o, E.A., Engelen, A.H., Renault, L., Garc\u00eda-March, J.R., Duarte, C.M., and Hendriks, I.E. (2018). Genetic and oceanographic tools reveal high population connectivity and diversity in the endangered pen shell Pinna nobilis. Sci. Rep., 8.","DOI":"10.1038\/s41598-018-23004-2"},{"key":"ref_77","doi-asserted-by":"crossref","unstructured":"Sanna, D., Azzena, I., Locci, C., Ankon, P., Kru\u017ei\u0107, P., Manfrin, C., Pallavicini, A., Ciriaco, S., Segarich, M., and Batistini, E. (2024). Reconstructing the evolutionary history of Pinna nobilis: New genetic signals from the past of a species on the brink of extinction. Animals, 14.","DOI":"10.3390\/ani14010114"},{"key":"ref_78","doi-asserted-by":"crossref","unstructured":"Peyran, C., Boissin, E., Morage, T., Nebot-Colomer, E., Iwankow, G., and Planes, S. (2021). Genetic homogeneity of the critically endangered fan mussel, Pinna nobilis, throughout lagoons of the Gulf of Lion (North-Western Mediterranean Sea). Sci. Rep., 11.","DOI":"10.1038\/s41598-021-87493-4"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"e8482","DOI":"10.1002\/ece3.8482","article-title":"Investigating population dynamics from parentage analysis in the highly endangered fan mussel Pinna nobilis","volume":"12","author":"Peyran","year":"2022","journal-title":"Ecol. Evol."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"e16491","DOI":"10.7717\/peerj.16491","article-title":"Insights on Pinna nobilis population genetic structure in the Aegean and Ionian Sea","volume":"11","author":"Sarafidou","year":"2023","journal-title":"PeerJ"},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"416","DOI":"10.12681\/mms.459","article-title":"New mitochondrial and nuclear primers for the Mediterranean marine bivalve Pinna nobilis","volume":"15","author":"Sanna","year":"2014","journal-title":"Mediterr. Mar. Sci."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"104795","DOI":"10.1016\/j.marenvres.2019.104795","article-title":"Age and growth of the endangered fan mussel Pinna nobilis in the western Mediterranean Sea","volume":"153","author":"Hernandis","year":"2020","journal-title":"Mar. Environ. Res."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"669","DOI":"10.12681\/mms.27137","article-title":"The last fortress fell: Mass mortality of Pinna nobilis in the Sea of Marmara","volume":"22","author":"Cinar","year":"2021","journal-title":"Mediterr. Mar. Sci."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"107265","DOI":"10.1016\/j.ocecoaman.2024.107265","article-title":"Looking for the surviving optimal areas for the threatened species Pinna nobilis in a highly anthropized coastal lagoon (Mar Menor, SE Spain)","volume":"255","year":"2024","journal-title":"Ocean Coast. Manag."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"21","DOI":"10.3354\/esr01357","article-title":"Population status of the Critically Endangered fan mussel Pinna nobilis in Thau Lagoon (France)","volume":"55","author":"Vicente","year":"2024","journal-title":"Endanger. Species Res."},{"key":"ref_86","doi-asserted-by":"crossref","unstructured":"Silvestri, S., Capra, V., Cucchiaro, S., Pivato, M., and Tarolli, P. (2022). Tides, topography, and seagrass cover controls on the spatial distribution of Pinna nobilis on a coastal lagoon tidal flat. J. Geophys. Res. Biogeosci., 127.","DOI":"10.1029\/2021JG006667"},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"126690","DOI":"10.1016\/j.jnc.2024.126690","article-title":"Lake faro (Central Mediterranean): A potential short-term reservoir for Pinna nobilis","volume":"81","author":"Lunetta","year":"2024","journal-title":"J. Nat. Conserv."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"107536","DOI":"10.1016\/j.ecss.2021.107536","article-title":"Noble fan-shell, Pinna nobilis, in Lake Faro (Sicily, Italy): Ineluctable decline or extreme opportunity?","volume":"261","author":"Donato","year":"2021","journal-title":"Estuar. Coast. Shelf Sci."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"679","DOI":"10.12681\/mms.35384","article-title":"Status, distribution, and threats of the last surviving fan mussel populations in Greece","volume":"24","author":"Papadakis","year":"2023","journal-title":"Mediterr. Mar. Sci."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"747","DOI":"10.12681\/mms.38613","article-title":"Pinna nobilis refugia breached: Ongoing Mass Mortality Event in the Gulf of Kalloni (Aegean Sea)","volume":"25","author":"Nikolaou","year":"2024","journal-title":"Mediterr. Mar. Sci."},{"key":"ref_91","unstructured":"DellaSala, D.A., and Goldstein, M.I. (2022). The Fan Mussel Pinna nobilis on the Brink of Extinction in the Mediterranean. Imperiled: The Encyclopedia of Conservation, Elsevier."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"105220","DOI":"10.1016\/j.marenvres.2020.105220","article-title":"Pinna nobilis in suboptimal environments are more tolerant to disease but more vulnerable to severe weather phenomena","volume":"163","author":"Prado","year":"2021","journal-title":"Mar. Environ. Res."},{"key":"ref_93","unstructured":"Barbier, E.B., Acreman, M., and Knowler, D. (1997). Economic Valuation of Wetlands: A Guide for Policy Makers and Planners, Ramsar Convention Bureau."},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"107","DOI":"10.1016\/j.marpolbul.2005.09.014","article-title":"Changes in benthic fish assemblages as a consequence of coastal works in a coastal lagoon: The Mar Menor (Spain, Western Mediterranean)","volume":"53","author":"Barcala","year":"2006","journal-title":"Mar. Pollut. Bull."},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"963","DOI":"10.1111\/j.1529-8817.2006.00261.x","article-title":"Positive feedback and the development and persistence of ecosystem disruptive algal blooms","volume":"42","author":"Sunda","year":"2006","journal-title":"J. Phycol."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1002\/aqc.3738","article-title":"Living under threat: Will one of the last Pinna nobilis populations be able to survive?","volume":"32","author":"Belando","year":"2022","journal-title":"Aquat. Conserv. Mar. Freshw. Ecosyst."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"114376","DOI":"10.1016\/j.marpolbul.2022.114376","article-title":"The relict population of Pinna nobilis in the Mar Menor is facing an uncertain future","volume":"185","author":"Murcia","year":"2022","journal-title":"Mar. Pollut. Bull."},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"378","DOI":"10.2307\/2405784","article-title":"Hybridization as an evolutionary stimulus","volume":"8","author":"Anderson","year":"1954","journal-title":"Evolution"},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"325","DOI":"10.1086\/674899","article-title":"Species collapse via hybridization in Darwin\u2019s tree finches","volume":"183","author":"Kleindorfer","year":"2014","journal-title":"Am. Nat."},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"229","DOI":"10.1111\/j.1420-9101.2012.02599.x","article-title":"Hybridization and speciation","volume":"26","author":"Abbott","year":"2013","journal-title":"J. Evol. Biol."},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"1055","DOI":"10.1111\/evo.12019","article-title":"Hybrid speciation and independent evolution in lineages of alpine butterflies","volume":"67","author":"Nice","year":"2013","journal-title":"Evolution"},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"923","DOI":"10.1038\/s41467-019-08809-7","article-title":"Hybridization is a recurrent evolutionary stimulus in wild yeast speciation","volume":"10","author":"Eberlein","year":"2019","journal-title":"Nat. Commun."},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"229","DOI":"10.1016\/j.tree.2005.02.010","article-title":"Hybridization as an invasion of the genome","volume":"20","author":"Mallet","year":"2005","journal-title":"Trends Ecol. Evol."},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"997","DOI":"10.1007\/s11033-020-06063-5","article-title":"Natural hybridization between pen shell species: Pinna rudis and the critically endangered Pinna nobilis may explain parasite resistance in P. nobilis","volume":"48","author":"Deudero","year":"2021","journal-title":"Mol. Biol. Rep."},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"e70227","DOI":"10.1002\/ece3.70227","article-title":"Internal transcribed spacer as effective molecular marker for the detection of natural hybridization between the bivalves Pinna nobilis and Pinna rudis","volume":"14","author":"Catanese","year":"2024","journal-title":"Ecol. Evol."},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"e10383","DOI":"10.1002\/ece3.10383","article-title":"The characterization of toll-like receptor repertoire in Pinna nobilis after mass mortality events suggests adaptive introgression","volume":"13","author":"Giantsis","year":"2023","journal-title":"Ecol. Evol."},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"109207","DOI":"10.1016\/j.ecolmodel.2020.109207","article-title":"Dynamic energy budget of endemic and critically endangered bivalve Pinna nobilis: A mechanistic model for informed conservation","volume":"434","author":"Haberle","year":"2020","journal-title":"Ecol. Model."},{"key":"ref_108","doi-asserted-by":"crossref","unstructured":"Kersting, D.K., V\u00e1zquez-Luis, M., Mourre, B., Belkhamssa, F.Z., \u00c1lvarez, E., Bakran-Petricioli, T., Barber\u00e1, C., Barraj\u00f3n, A., Cort\u00e9s, E., and Deudero, S. (2020). Recruitment disruption and the role of unaffected populations for potential recovery after the Pinna nobilis mass mortality event. Front. Mar. Sci., 7.","DOI":"10.3389\/fmars.2020.594378"},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"1501","DOI":"10.1002\/aqc.4009","article-title":"Reproduction of the endangered fan mussel Pinna nobilis under natural and captivity conditions","volume":"33","author":"Hernandis","year":"2023","journal-title":"Aquat. Conserv. Mar. Freshw. Ecosyst."},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"e70565","DOI":"10.1002\/ece3.70565","article-title":"Handling of the Bivalve Pinna nobilis, Endangered and Pathogen-Affected Species, for Controlled Reproduction: Precautions Taken","volume":"14","author":"Ferranti","year":"2024","journal-title":"Ecol. Evol."},{"key":"ref_111","doi-asserted-by":"crossref","first-page":"141","DOI":"10.1006\/jipa.2001.5052","article-title":"Epizootiology and pathology of juvenile oyster disease in the eastern oyster, Crassostrea virginica","volume":"78","author":"Ford","year":"2001","journal-title":"J. Invertebr. Pathol."},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"735167","DOI":"10.1016\/j.aquaculture.2020.735167","article-title":"Growth of juvenile Pinna nobilis in captivity conditions: Dietary and pathological constraints","volume":"522","author":"Prado","year":"2020","journal-title":"Aquaculture"},{"key":"ref_113","unstructured":"Hernandis Caballero, S. (2021). Avances en la Ecolog\u00eda y Biolog\u00eda Para la Conservaci\u00f3n de la Especie Cr\u00edticamente Amenazada Pinna nobilis (Linnaeus, 1758), Endemismo Mediterr\u00e1neo. [Ph.D. Thesis, Universidad Cat\u00f3lica de Valencia]."},{"key":"ref_114","doi-asserted-by":"crossref","unstructured":"Hernandis, S., Ibarrola, I., Tena-Medialdea, J., V\u00e1zquez-Luis, M., Garc\u00eda-March, J.R., Prado, P., and Albentosa, M. (2022). Scope for growth and dietary needs of Mediteranean Pinnids maintained in captivity. BMC Zool., 7.","DOI":"10.1186\/s40850-022-00141-w"},{"key":"ref_115","doi-asserted-by":"crossref","first-page":"350","DOI":"10.33714\/masteb.627562","article-title":"Population, aquaculture and transplantation applications of critically endangered species Pinna nobilis (Linnaeus 1758) in the Mediterranean Sea","volume":"10","author":"Acarli","year":"2021","journal-title":"Mar. Sci. Technol. Bull."},{"key":"ref_116","doi-asserted-by":"crossref","first-page":"1371","DOI":"10.1111\/raq.12654","article-title":"Worldwide overview of reproduction, juvenile collection, spat production and cultivation of pen shells","volume":"14","year":"2022","journal-title":"Rev. Aquac."},{"key":"ref_117","first-page":"55","article-title":"Protocole pour la transplantation des nacres Pinna nobilis dans divers substrats","volume":"18","author":"Trigos","year":"2016","journal-title":"Mar. Life"},{"key":"ref_118","doi-asserted-by":"crossref","first-page":"736964","DOI":"10.1016\/j.aquaculture.2021.736964","article-title":"Suspended culture of Pinna rudis enhances survival and allows the development of a seasonal growth model for Mediterranean Pinnids","volume":"543","author":"Hernandis","year":"2021","journal-title":"Aquaculture"},{"key":"ref_119","first-page":"77","article-title":"Phytoplankton profitability and use as organic matter source by Pinna nobilis","volume":"7","author":"Trigos","year":"2015","journal-title":"Nereis"},{"key":"ref_120","doi-asserted-by":"crossref","first-page":"102","DOI":"10.1016\/j.aquaculture.2017.10.015","article-title":"Adult spawning and early larval development of the endangered bivalve Pinna nobilis","volume":"483","author":"Trigos","year":"2018","journal-title":"Aquaculture"},{"key":"ref_121","doi-asserted-by":"crossref","first-page":"247","DOI":"10.1016\/S0006-3207(97)00081-5","article-title":"Flagships, umbrellas, and keystones: Is single-species conservation effective in forest landscapes?","volume":"83","author":"Simberloff","year":"1998","journal-title":"Biol. Conserv."},{"key":"ref_122","doi-asserted-by":"crossref","first-page":"76","DOI":"10.1111\/j.1523-1739.2004.00450.x","article-title":"Usefulness of the umbrella species concept for conservation planning: A case study with the white-backed woodpecker","volume":"18","author":"Roberge","year":"2004","journal-title":"Biol. Conserv."}],"container-title":["Animals"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2076-2615\/15\/14\/2044\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,9]],"date-time":"2025-10-09T18:08:27Z","timestamp":1760033307000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2076-2615\/15\/14\/2044"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,7,11]]},"references-count":122,"journal-issue":{"issue":"14","published-online":{"date-parts":[[2025,7]]}},"alternative-id":["ani15142044"],"URL":"https:\/\/doi.org\/10.3390\/ani15142044","relation":{},"ISSN":["2076-2615"],"issn-type":[{"value":"2076-2615","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,7,11]]}}}