Climate-driven habitat shifts of the vulnerable sea pen Funiculina quadrangularis in the Mediterranean Sea

Open Access

Issue		Aquat. Living Resour. Volume 39, 2026


Article Number		11
Number of page(s)		30
DOI		https://doi.org/10.1051/alr/2026004
Published online		28 April 2026

AdriaMed. 2011. Report of the Technical meeting on SoleMon survey activities, May 2011. GCP/RER/010/ITA/SR-01. (No. 01), AdriaMed Scientific Reports. FAO, Rome. [Google Scholar]
Aguilar R, Perry AL, López J, Conservation and management of vulnerable marine benthic ecosystems, in: Rossi, S., Bramanti, L., Gori, A., Orejas, C. (Eds.), Marine Animal Forests: The Ecology of Benthic Biodiversity Hotspots. Springer International Publishing, Cham, 2017, pp. 1–43. https://doi.org/10.1007/978-3-319-17001-5_34-1 [Google Scholar]
Allouche O, Tsoar A, Kadmon R. 2006. Assessing the accuracy of species distribution models: prevalence, kappa and the true skill statistic (TSS). J Appl Ecol 43: 1223–1232. [Google Scholar]
Ambroso S, Dominguez-Carrió C, Grinyó J, López-González PJ, Gili JM, Purroy A, Requena S, Madurell T. 2013. In situ observations on withdrawal behaviour of the sea pen Virgularia mirabilis. Mar Biodivers 43: 257–258. [Google Scholar]
Anderson OF, Stephenson F, Behrens E, Rowden AA. 2022. Predicting the effects of climate change on deep-water coral distribution around New Zealand–will there be suitable refuges for protection at the end of the 21st century? Glob Change Biol 28: 6556–6576. [Google Scholar]
Araújo MB, Guisan A. 2006. Five (or so) challenges for species distribution modelling. J Biogeogr 33: 1677–1688. [Google Scholar]
Arrêté du 21 octobre 2024. https://www.legifrance.gouv.fr/jorf/id/JORFTEXT000050397265 [Google Scholar]
Ardizzone GD. 2006. Sensitive and Essential Fish Habitats in the Mediterranean Sea. Working document to the STECF/SGMED-06-01 sub-group meeting on sensitive and essential fish habitats in the Mediterranean. https://stecf.jrc.ec.europa.eu/documents/d/stecf/plen_06-01 [Google Scholar]
Assis J, Fernández Bejarano SJ, Salazar VW, Schepers L, Gouvêa L, Fragkopoulou E, Leclercq F, Vanhoorne B, Tyberghein L, Serrão EA, Verbruggen H, De Clerck O. 2024. Bio-ORACLE v3.0: pushing marine data layers to the CMIP6 Earth system models of climate change research. Glob Ecol Biogeogr 33: e13813. [Google Scholar]
Augier H. Inventory and classification of marine benthic biocenoses of the Mediterranean, in: Council of Europe (Ed.), Nature and Environment Series, Vol. 25, Council of Europe, Strasbourg: Croton, N.Y., 1982 [Google Scholar]
Austin MP. 2002. Spatial prediction of species distribution: an interface between ecological theory and statistical modelling. Ecol Model 157: 101–118. [Google Scholar]
Baillon S, Hamel JF, Mercier A. 2014. Diversity, distribution and nature of faunal associations with deep-sea pennatulacean corals in the northwest Atlantic. PLoS One 9: e111519. [Google Scholar]
Bastari A, Pica D, Ferretti F, Micheli F, Cerrano C. 2018. Sea pens in the Mediterranean Sea: habitat suitability and opportunities for ecosystem recovery. ICES J Mar Sci 75: 1722–1732. [Google Scholar]
Bateman DC, Bishop MJ. 2017. The environmental context and traits of habitat-forming bivalves influence the magnitude of their ecosystem engineering. Mar Ecol Prog Ser 563: 95–110. https://doi.org/10.3354/meps11959 [Google Scholar]
Beazley L, Wang Z, Kenchington E, Yashayaev I, Rapp HT, Xavier JR, Murillo FJ, Fenton D, Fuller S. 2018. Predicted distribution of the glass sponge Vazella pourtalesi on the Scotian Shelf and its persistence in the face of climatic variability. PLoS One 13: e0205505. https://doi.org/10.1371/journal.pone.0205505 [Google Scholar]
Beazley LI, Kenchington EL, Murillo FJ, Sacau MM. 2013. Deep-sea sponge grounds enhance diversity and abundance of epibenthic megafauna in the northwest Atlantic. ICES J Mar Sci 70: 1471–1490. https://doi.org/10.1093/icesjms/fst124 [Google Scholar]
Bo M, Otero M. del M, Numa C. 2017. Overview of the conservation status of Mediterranean anthozoa. IUCN. https://doi.org/10.2305/IUCN.CH.2017.RA.2.en [Google Scholar]
Bolam SG, Fernandes TF. 2003. Dense aggregations of Pygospio elegans: effect on macrofaunal community structure and sediments. J Sea Res 49: 171–185. https://doi.org/10.1016/S1385-1101(03)00007-8 [Google Scholar]
Bourque JR, Demopoulos AWJ. 2018. The influence of different deep-sea coral habitats on sediment macrofaunal community structure and function. PeerJ 6: e5276. https://doi.org/10.7717/peerj.5276 [Google Scholar]
Braunisch V, Coppes J, Arlettaz R, Suchant R, Schmid H, Bollmann K. 2013. Selecting from correlated climate variables: a major source of uncertainty for predicting species distributions under climate change. Ecography 36: 971–983. https://doi.org/10.1111/j.1600-0587.2013.00138.x [Google Scholar]
Breiman L. 2001. Random Forests. Mach Learn 45: 5–32. https://doi.org/10.1023/A:1010933404324 [Google Scholar]
Buhl-Mortensen L, Vanreusel A, Gooday AJ, Levin LA, Priede IG, Buhl-Mortensen P, Gheerardyn H, King NJ, Raes M. 2010. Biological structures as a source of habitat heterogeneity and biodiversity on the deep ocean margins. Mar Ecol 31: 21–50. https://doi.org/10.1111/j.1439-0485.2010.00359.x [Google Scholar]
Byrne RH, Mecking S, Feely RA, Liu X. 2010. Direct observations of basin-wide acidification of the North Pacific Ocean. Geophys Res Lett 37: L02601. https://doi.org/10.1029/2009GL040999 [Google Scholar]
Carbonara P, Zupa W, Follesa MC, Cau A, Capezzuto F, Chimienti G, DOnghia G, Lembo G, Pesci P, Porcu C, Bitetto I, Spedicato MT, Maiorano P. 2020. Exploring a deep-sea vulnerable marine ecosystem: Isidella elongata species assemblages in the western and central Mediterranean. Deep Sea Res Part I Oceanogr Res Pap 166: 103406. https://doi.org/10.1016/j.dsr.2020.103406 [Google Scholar]
Carbonara P, Zupa W, Follesa MC, Cau A, Donnaloia M, Alfonso S, Casciaro L, Spedicato MT, Maiorano P. 2022. Spatio-temporal distribution of Isidella elongata in the southern Adriatic Sea. Hydrobiologia 849: 4837–4855. https://doi.org/10.1007/s10750-022-05022-4 [Google Scholar]
Carter AL, Wilson AMW, Bello M, Hoyos-Padilla EM, Inall ME, Ketchum JT, Schurer A, Tudhope AW. 2020. Assessing opportunities to support coral reef climate change refugia in MPAs. Mar Policy 112: 103769. https://doi.org/10.1016/j.marpol.2019.103769 [Google Scholar]
Cartes JE, Maynou F, Sardà F, Company JB, Lloris D, Tudela S. 2004. Mediterranean deep-sea ecosystems: an overview of their diversity, structure, functioning and anthropogenic impacts, in: The Mediterranean Deep-Sea Ecosystems: An Overview of Their Diversity, Structure, Functioning and Anthropogenic Impacts, with a Proposal for Conservation, IUCN, Málaga and WWF, Rome, 2004, pp. 9–38. [Google Scholar]
Certain G, Jadaud A, Vaz S, Metral L, Cheret I, Billet N, Schull, Tessier E, Cornou AS, Souplet A, 2025. MEDITS catch data: bentho-demersal assemblages of the Gulf of Lion and eastern Corsica (North-western Mediterranean) from 2016 to present. SEANOE. https://doi.org/10.17882/99052 [Google Scholar]
Chimienti G, Angeletti L, Mastrototaro F. 2018a. Withdrawal behaviour of the red sea pen Pennatula rubra. Eur Zool J 85: 64–70. https://doi.org/10.1080/24750263.2018.1438530 [Google Scholar]
Chimienti G, Angeletti L, Rizzo L, Tursi A, Mastrototaro F. 2018b. ROV vs trawling approaches in the study of benthic communities. J Mar Biol Assoc UK 98: 1859–1869. https://doi.org/10.1017/S0025315418000851 [Google Scholar]
Chollett I, Escovar-Fadul X, Schill SR, Croquer A, Dixon AM, Beger M, Shaver E, Pietsch McNulty V, Wolff NH. 2022. Planning for resilience: incorporating uncertainty when prioritizing climate refugia. Glob Change Biol 28: 4054–4068. https://doi.org/10.1111/gcb.16167 [Google Scholar]
Cliff AD, Ord K. 1970. Spatial autocorrelation: a review of existing and new measures with applications. Econ Geogr 46: 269–292. https://doi.org/10.2307/143144 [Google Scholar]
Colloca F, Carpentieri P, Balestri E, Ardizzone GD. 2004. A critical habitat for Mediterranean fish resources. Mar Biol 145: 1129–1142. https://doi.org/10.1007/s00227-004-1405-8 [Google Scholar]
Combes M, Vaz S, Grehan A, Morato T, Arnaud-Haond S, Dominguez-Carrió C, Fox A, González-Irusta JM, Johnson D, Callery O, Davies A, Fauconnet L, Kenchington E, Orejas C, Roberts JM, Taranto G, Menot L. 2021. Systematic conservation planning at an ocean basin scale. Front Mar Sci 8. https://doi.org/10.3389/fmars.2021.611358 [Google Scholar]
Danovaro R. 2018. Climate change impacts on the biota of the deep Mediterranean Sea. Rend Lincei Sci Fis Nat 29: 525–541. https://doi.org/10.1007/s12210-018-0725-4 [Google Scholar]
Davies AJ, Guinotte JM. 2011. Global habitat suitability for framework-forming cold-water corals. PLoS One 6: e18483. https://doi.org/10.1371/journal.pone.0018483 [Google Scholar]
De Clippele LH, Buhl-Mortensen P, Buhl-Mortensen L. 2015. Fauna associated with cold-water gorgonians and sea pens. Cont Shelf Res 105: 67–78. https://doi.org/10.1016/j.csr.2015. 06.007 [Google Scholar]
Diniz-Filho JAF, Bini LM, Rangel TF, Loyola RD, Hof C, Nogués-Bravo D, Araújo MB. 2009. Partitioning and mapping uncertainties in ensembles of forecasts. Ecography 32: 897–906. https://doi.org/10.1111/j.1600-0587.2009.06196.x [Google Scholar]
Dormann CF, Elith J, Bacher S, Buchmann C, Carl G, Carré G, Marquéz JRG, Gruber B, Lafourcade B, Leitão PJ, Münkemüller T, McClean C, Osborne PE, Reineking B, Schröder B, Skidmore AK, Zurell D, Lautenbach S. 2013. Collinearity: a review of methods to deal with it. Ecography 36: 27–46. https://doi.org/10.1111/j.1600-0587.2012.07348.x [Google Scholar]
Dormann CF, McPherson JM, Araújo MB, Bivand R, Bolliger J, Carl G, Davies RG, Hirzel A, Jetz W, Kissling WD, Kühn I, Ohlemüller R, Peres-Neto PR, Reineking B, Schröder B, Schurr FM, Wilson R. 2007. Methods to account for spatial autocorrelation in species distribution data. Ecography 30: 609–628. https://doi.org/10.1111/j.2007.0906-7590.05171.x [Google Scholar]
Downie AL, Vieira RP, Hogg OT, Darby C. 2021. Distribution of vulnerable marine ecosystems at the South Sandwich Islands: results from the Blue Belt Discovery Expedition 99 deep-water camera surveys. Front Mar Sci 8: 662285. https://doi.org/10.3389/fmars.2021.662285 [Google Scholar]
Duffy JE. 2006. Biodiversity and the functioning of seagrass ecosystems. Mar Ecol Prog Ser 311: 233–250. https://doi.org/10.3354/meps311233 [Google Scholar]
Edwards DCB, Moore CG. 2009. Reproduction in the sea pen Funiculina quadrangularis (Anthozoa: Pennatulacea) from the west coast of Scotland. Estuar Coast Shelf Sci 82: 161–168. https://doi.org/10.1016/j.ecss.2009.01.006 [Google Scholar]
Eigaard OR, Bastardie F, Hintzen NT, Buhl-Mortensen L, Buhl-Mortensen P, Catarino R, Dinesen GE, Egekvist J, Fock HO, Geitner K, Gerritsen HD, González MM, Jonsson P, Kavadas S, Laffargue P, Lundy M, Gonzalez-Mirelis G, Nielsen JR, Papadopoulou N, Posen PE, Pulcinella J, Russo T, Sala A, Silva C, Smith CJ, Vanelslander B, Rijnsdorp AD. 2017. The footprint of bottom trawling in European waters: distribution, intensity, and seabed integrity. ICES J Mar Sci 74: 847–865. https://doi.org/10.1093/icesjms/fsw194 [Google Scholar]
Elith J, Leathwick JR. 2009. Species distribution models: ecological explanation and prediction across space and time. Annu Rev Ecol Evol Syst 40: 677–697. https://doi.org/10.1146/annurev.ecolsys.110308.120159 [Google Scholar]
Elith J, Leathwick JR, Hastie T. 2008. A working guide to boosted regression trees. J Anim Ecol 77: 802–813. https://doi.org/10.1111/j.1365-2656.2008.01390.x [Google Scholar]
Emig CC. 1997. Bathyal zones of the Mediterranean continental slope: an attempt. Publ Espec Inst Esp Oceanogr 23: 23–33. [Google Scholar]
Eno N. 2001. Effects of crustacean traps on benthic fauna. ICES J Mar Sci 58: 11–20. https://doi.org/10.1006/jmsc.2000.0984 [Google Scholar]
Fabri M-C, Pedel L, Beuck L, Galgani F, Hebbeln D, Freiwald A. 2014. Megafauna of vulnerable marine ecosystems in French Mediterranean submarine canyons: spatial distribution and anthropogenic impacts. Deep Sea Res Part II Top Stud Oceanogr 104: 184–207. https://doi.org/10.1016/j.dsr2.2013.06.016 [Google Scholar]
Fabri MC, Pedel L. 2020. Funiculina quadrangularis (Pennatulacea) presence and absence extracted from video, from Fabri et al 2014. Sextant. https://doi.org/10.1016/j.dsr2.2013.06.016 [Google Scholar]
FAO. 2009. International Guidelines for the Management of Deep-sea Fisheries in the High Seas/Directives internationales sur la gestion de la pêche profonde en haute mer/ Directrices Internacionales para la Ordenación de las Pesquerías de Aguas Profundas en Alta Mar. FAO, Rome 14: 20. https://www.fao.org/documents/card/en/c/b02fc35e-a0c4-545a-86fb-4fc340e13b52 [Google Scholar]
Farriols MT, Irlinger C, Ordines F, Palomino D, Marco-Herrero E, Soto-Navarro J, Jordà G, Mallol S, Díaz D, Martínez-Carreño N, Díaz JA, Fernandez-Arcaya U, Joher S, Ramírez-Amaro S, R de la Ballina N, Vázquez JT, Massutí E. 2022. Recovery signals of rhodolith beds since bottom trawling ban in the SCI Menorca Channel (Western Mediterranean). Diversity 14: 20. https://doi.org/10.3390/d14010020 [Google Scholar]
Fawcett T. 2006. An introduction to ROC analysis. Pattern Recognit Lett 27: 861–874. https://doi.org/10.1016/j.patrec.2005.10.010 [Google Scholar]
Fourcade Y, Besnard AG, Secondi J. 2018. Paintings predict the distribution of species, or the challenge of selecting environmental predictors and evaluation statistics. Glob Ecol Biogeogr 27: 245–256. https://doi.org/10.1111/geb.12684 [Google Scholar]
Fujita T, Ohta S. 1988. Photographic observations of the life style of a deep-sea ophiuroid Asteronyx loveni (Echinodermata). Deep Sea Res Part A Oceanogr Res Pap 35: 2029–2043. https://doi.org/10.1016/0198-0149(88)90123-9 [Google Scholar]
Fourt M, Goujard A, Perez T, Chevaldonné P. 2017. Guide de la faune profonde de la mer Méditerranée. Exploration des roches et canyons sous-marins des côtes françaises. Muséum national d’Histoire naturelle. Publications scientifiques, Paris. http://sciencepress.mnhn.fr/fr/collections/patrimoines-naturels/guide-de-la-faune-profonde-de-la-mer-mediterranee [Google Scholar]
Georges V, Vaz S, Carbonara P, Fabri MC, Fanelli E, Follesa MC, Garofalo G, Gerovasileiou V, Jadaud A, Maiorano P, Marin P, Mytilineou C, Orejas C, Del Mar Otero M, Smith CJ, Thasitis I, Lauria V. 2024. Mapping the habitat refugia of Isidella elongata under climate change and trawling impacts to preserve vulnerable marine ecosystems in the Mediterranean. Sci Rep 14: 6246. https://doi.org/10.1038/s41598-024-56338-1 [Google Scholar]
Georgian SE, Anderson OF, Rowden AA. 2019. Ensemble habitat suitability modeling of vulnerable marine ecosystem indicator taxa to inform deep-sea fisheries management in the South Pacific Ocean. Fish Res 211: 256–274. https://doi.org/10.1016/j.fishres.2018.11.020 [Google Scholar]
GFCM SAC, 2009. Criteria for the identification of sensitive habitats of relevance for the management of priority species. General Fisheries Commission for the Mediterranean (GFCM), Scientific Advisory Committee (SAC), Sub-Committee on Marine Environment and Ecosystems, Malaga. [Google Scholar]
Giangrande A, Gambi MC, Gravina MF. 2020. Polychaetes as habitat former: structure and function, in: Rossi S, Bramanti L (Eds.), Perspectives on the Marine Animal Forests of the World. Springer, Cham, 2020, pp. 219–237. https://doi.org/10.1007/978-3-030-57054-5_8 [Google Scholar]
Gori A, Ferrier-Pagès C, Hennige SJ, Murray F, Rottier C, Wicks LC, Roberts JM. 2016. Physiological response of the cold-water coral Desmophyllum dianthus to thermal stress and ocean acidification. Peer J 4: e1606. https://doi.org/10.7717/peerj.1606 [Google Scholar]
Greathead CF, Donnan DW, Mair JM, Saunders GR. 2007. The sea pens Virgularia mirabilis, Pennatula phosphorea and Funiculina quadrangularis: distribution and conservation issues in Scottish waters. J Mar Biol Assoc UK 87: 1095–1103. https://doi.org/10.1017/S0025315407056238 [Google Scholar]
Greathead C, González-Irusta JM, Clarke J, Boulcott P, Blackadder L, Weetman A, Wright PJ. 2015. Environmental requirements for three sea pen species: relevance to distribution and conservation. ICES J Mar Sci 72: 576–586. https://doi.org/10.1093/icesjms/fsu129 [Google Scholar]
Greeley K. 2022. Skeletal growth ring microanalysis and growth rates in sea pens from the Laurentian Channel MPA, Newfoundland. Memorial Univ Newfoundland. https://doi.org/10.48336/t646-sy04 [Google Scholar]
Grinyó J, Gori A, Greenacre M, Requena S, Canepa A, Lo Iacono C, Ambroso S, Purroy A, Gili JM. 2018. Megabenthic assemblages in the continental shelf edge and upper slope of the Menorca Channel, Western Mediterranean Sea. Prog Oceanogr 162: 40–51. https://doi.org/10.1016/j.pocean.2018.02.002 [Google Scholar]
Hastie T, Tibshirani R. 1986. Generalized additive models. Stat Sci 1: 297–310. https://doi.org/10.1214/ss/1177013604 [Google Scholar]
Hattab T, Garzón-López CX, Ewald M, Skowronek S, Aerts R, Horen H, Brasseur B, Gallet-Moron E, Spicher F, Decocq G, Feilhauer H, Honnay O, Kempeneers P, Schmidtlein S, Somers B, Van De Kerchove R, Rocchini D, Lenoir J. 2017. A unified framework to model the potential and realized distributions of invasive species within the invaded range. Divers Distrib 23: 806–819. https://doi.org/10.1111/ddi.12566 [Google Scholar]
Hughes DJ. 1998. Sea pens and burrowing megafauna. An overview of dynamics and sensitivity characteristics for conservation management of marine SACs. Scottish Assoc Mar Sci (UK Marine SACs Project) 105. http://ukmpa.marinebiodiversity.org/uk_sacs/pdfs/seapens.pdf [Google Scholar]
Kaiser MJ, Collie JS, Hall SJ, Jennings S, Poiner IR. 2002. Modification of marine habitats by trawling activities: prognosis and solutions. Fish Fish 3: 114–136. https://doi.org/10.1046/j.1467-2979.2002.00079.x [Google Scholar]
Karp MA, Cimino M, Craig JK, Crear DP, Haak C, Hazen EL, Kaplan I, Kobayashi DR, Moustahfid H, Muhling B, Pinsky ML, Smith LA, Thorson JT, Woodworth-Jefcoats PA. 2025. Applications of species distribution modeling and future needs to support marine resource management. ICES J Mar Sci 82: fsaf024. https://doi.org/10.1093/icesjms/fsaf024 [Google Scholar]
Keeling RF, Körtzinger A, Gruber N. 2010. Ocean deoxygenation in a warming world. Annu Rev Mar Sci 2: 199–229. https://doi.org/10.1146/annurev.marine.010908.163855 [Google Scholar]
Keppel G, Mokany K, Wardell-Johnson GW, Phillips BL, Welbergen JA, Reside AE. 2015. The capacity of refugia for conservation planning under climate change. Front Ecol Environ 13: 106–112. https://doi.org/10.1890/140055 [Google Scholar]
Lauria V, Garofalo G, Fiorentino F, Massi D, Milisenda G, Piraino S, Russo T, Gristina M. 2017. Species distribution models of two critically endangered deep-sea octocorals reveal fishing impacts on vulnerable marine ecosystems in central Mediterranean Sea. Sci Rep 7: 8049. https://doi.org/10.1038/s41598-017-08386-z [Google Scholar]
Lauria V, Massi D, Fiorentino F, Milisenda G, Cillari T. 2021. Habitat suitability mapping of the black coral Leiopathes glaberrima to support conservation of vulnerable marine ecosystems. Sci Rep 11: 15661. https://doi.org/10.1038/s41598-021-95256-4 [Google Scholar]
Lauria V, Gargano F, Arculeo M, Bono G, Fiorentino F, Flot JF, Collard O, Follesa MC, Garofalo G, Georges V, Maiorano P, Massi D, Stamouli C, Vaz S, Vitale S, Vecchioni L. 2025. Genetic structure and connectivity of the critically endangered deep-sea octocoral Isidella elongata: Implications for marine biodiversity conservation in the Mediterranean Sea. Medit Mar Sci 26: 1050–1067. https://doi.org/10.12681/mms.40883 [Google Scholar]
Legendre P,& Legendre L. Numerical Ecology, 3rd ed, Elsevier, 1998. [Google Scholar]
Lejeusne C, Chevaldonné P, Pergent-Martini C, Boudouresque CF, Pérez T. 2010. Climate change effects on a miniature ocean: the highly diverse, highly impacted Mediterranean Sea. Trends Ecol Evol 25: 250–260. https://doi.org/10.1016/j.tree.2009.10.009 [Google Scholar]
Leroy B, Bellard C, Dubos N, Colliot A, Vasseur M, Courtial C, Bakkenes M, Canard A, Ysnel F. 2014. Forecasted climate and land use changes, and protected areas: the contrasting case of spiders. Divers Distrib 20: 686–697. https://doi.org/10.1111/ddi.12191 [Google Scholar]
Macias D, Bisselink B, Carmona-Moreno C, Druon JN, Duteil O, Garcia-Gorriz E, Grizzetti B, Guillen J, Miladinova S, Pistocchi A, Piroddi C, Polimene L, Serpetti N, Stips A, Trichakis I, Udias A, Vigiak O. 2025. The overlooked impacts of freshwater scarcity on oceans as evidenced by the Mediterranean Sea. Nat Commun 16: 998. https://doi.org/10.1038/s41467-024-54979-4 [Google Scholar]
Mačić V, Đorđević N, Đurović M, Petović S, Russo T. 2022. Improving knowledge of Funiculina quadrangularis and vulnerable marine ecosystems in the south Adriatic. Medit Mar Sci 23: 805–816. https://doi.org/10.12681/mms.29898 [Google Scholar]
Manea E, Bianchelli S, Fanelli E, Danovaro R, Gissi E. 2020. Towards an ecosystem-based marine spatial planning in the deep Mediterranean Sea. Sci Total Environ 715: 136884. https://doi.org/10.1016/j.scitotenv.2020.136884 [Google Scholar]
Maldonado M, Aguilar R, Bannister RJ, Bell JJ, Conway KW, Dayton PK, Díaz C, Gutt J, Kelly M, Kenchington ELR, Leys SP, Pomponi SA, Rapp HT, Rützler K, Tendal OS, Vacelet J, Young CM. Sponge grounds as key marine habitats: A synthetic review of types, structure, functional roles, and conservation concerns, in: Rossi S, Bramanti L, Gori A, Orejas Saco del Valle C (Eds.), Marine Animal Forests: The Ecology of Benthic Biodiversity Hotspots. Springer, Cham, 2015, pp. 1–39. https://doi.org/10.1007/978-3-319-17001-5_24-1 [Google Scholar]
Margirier F, Testor P, Heslop E, Mallil K, Bosse A, Houpert L, Mortier L, Bouin MN, Coppola L, D’Ortenzio F, Durrieu de Madron X, Mourre B, Prieur L, Raimbault P, Taillandier V. 2020. Abrupt warming and salinification of intermediate waters interplays with decline of deep convection in the Northwestern Mediterranean Sea. Sci Rep 10: 20923. https://doi.org/10.1038/s41598-020-77859-5 [Google Scholar]
Margules CR, Pressey RL, 2000. Systematic conservation planning. Nature 405, 243–253. https://doi.org/10.1038/35012251 [Google Scholar]
Marsaglia L, Parisi A, Libralato S, Miller NA, Davis P, Paolo FS, Fiorentino F, Garofalo G, Costantini M, Russo T. 2024. Shedding light on trawl fishing activity in the Mediterranean Sea with remote sensing data. ICES J Mar Sci 82: fsae153. https://doi.org/10.1093/icesjms/fsae153 [Google Scholar]
Martinelli M, Zacchetti L, Belardinelli A, Domenichetti F, Scarpini P, Penna P, Medvešek D, Isajlović I, Vrgoč N. 2023. Changes in abundance and distribution of the sea pen Funiculina quadrangularis in the central Adriatic Sea (Mediterranean Basin) in response to variations in trawling intensity. Fishes 8: 347. https://doi.org/10.3390/fishes8070347 [Google Scholar]
Matos FL, Aguzzi J, Company JB, Cunha MR. 2024. Gone with the stream: Functional connectivity of a cold-water coral at basin scale. Limnol Oceanogr 69: 217–231. https://doi.org/10.1002/lno.12444 [Google Scholar]
Maynou F, Cartes JE. 2012. Effects of trawling on fish and invertebrates from deep-sea coral facies of Isidella elongata in the western Mediterranean. J Mar Biol Assoc UK 92: 1501–1507. https://doi.org/10.1017/S0025315411001603 [Google Scholar]
McCullagh P. 1984. Generalized linear models. Eur J Oper Res 16: 285–292. https://doi.org/10.1016/0377-2217(84)90282-0 [Google Scholar]
MEDITS Working Group. 2017. MEDITS Handbook. Version 9, 106. https://www.sibm.it/MEDITS%202011/docs/Medits_Handbook_2017_version_9_5-60417r.pdf [Google Scholar]
Mesgaran MB, Cousens RD, Webber BL. 2014. Here be dragons: a tool for quantifying novelty due to covariate range and correlation change when projecting species distribution models. Divers Distrib 20: 1147–1159. https://doi.org/10.1111/ddi.12209 [Google Scholar]
Millot J, Georges V, Lauria V, Hattab T, Dominguez-Carrió C, Gerovasileiou V, Smith CJ, Mytilineou C, Teresa Farriols M, Fabri MC, Carbonara P, Massi D, Rinelli P, Profeta A, Chimienti G, Jadaud A, Thasitis I, Camilleri K, Mifsud J, Vaz S. 2024. Habitat shifts of the vulnerable crinoid Leptometra phalangium under climate change scenarios. Prog Oceanogr 229: 103355. https://doi.org/10.1016/j.pocean.2024.103355 [Google Scholar]
Millot J, Vaz S. 2024. Modelled seabed sediment average grain size in the European North-East Atlantic and Mediterranean waters. Sextant. https://doi.org/10.12770/79050557-6f8a-48cc-9e60-4112cae9fd18 [Google Scholar]
Morato T, González-Irusta JM, Dominguez-Carrió C, Wei CL, Davies A, Sweetman AK, Taranto GH, Beazley L, García-Alegre A, Grehan A, Laffargue P, Murillo FJ, Sacau M, Vaz S, Kenchington E, Arnaud-Haond S, Callery O, Chimienti G, Cordes E, Egilsdottir H, Freiwald A, Gasbarro R, Gutiérrez-Zárate C, Gianni M, Gilkinson K, Wareham Hayes VE, Hebbeln D, Hedges K, Henry LA, Johnson D, Koen-Alonso M, Lirette C, Mastrototaro F, Menot L, Molodtsova T, Durán Muñoz P, Orejas C, Pennino MG, Puerta P, Ragnarsson SÁ, Ramiro-Sánchez B, Rice J, Rivera J, Roberts JM, Ross SW, Rueda JL, Sampaio Í, Snelgrove P, Stirling D, Treble MA, Urra J, Vad J, van Oevelen D, Watling L, Walkusz W, Wienberg C, Woillez M, Levin LA, Carreiro-Silva M. 2020. Climate-induced changes in the suitable habitat of cold-water corals and commercially important deep-sea fishes in the North Atlantic. Glob Change Biol 26: 2181–2202. https://doi.org/10.1111/gcb.14996 [Google Scholar]
Murillo F, Kenchington E, Tompkins G, Beazley L, Baker E, Knudby A, Walkusz W. 2018. Sponge assemblages and predicted archetypes in the eastern Canadian Arctic. Mar Ecol Prog Ser 597: 115–135. https://doi.org/10.3354/meps12589 [Google Scholar]
Owens HL, Campbell LP, Dornak LL, Saupe EE, Barve N, Soberón J, Ingenloff K, Lira-Noriega A, Hensz CM, Myers CE, Peterson AT. 2013. Constraints on interpretation of ecological niche models by limited environmental ranges on calibration areas. Ecol Model 263: 10–18. https://doi.org/10.1016/j.ecolmodel.2013.04.011 [Google Scholar]
Pastor F, Valiente JA, Khodayar S. 2020. A warming Mediterranean: 38 years of increasing sea surface temperature. Remote Sens 12: 2687. https://doi.org/10.3390/rs12172687 [Google Scholar]
Pellissier L, Bråthen KA, Vittoz P, Yoccoz NG, Dubuis A, Meier ES, Zimmermann NE, Randin CF, Thuiller W, Garraud L, Van Es J, Guisan A. 2013. Thermal niches are more conserved at cold than warm limits in arctic-alpine plant species. Glob Ecol Biogeogr 22: 933–941. https://doi.org/10.1111/geb.12057 [Google Scholar]
Pérès JM, Picard J. Nouveau Manuel de bionomie benthique de la mer Méditerranée, in: Recueil Des Travaux de La Station Marine d’Endoume, 1964, p. 137. [Google Scholar]
Peterson AT, Cobos ME, Jiménez-García D. 2018. Major challenges for correlational ecological niche model projections to future climate conditions. Ann NY Acad Sci 1429: 66–77. https://doi.org/10.1111/nyas.13873 [Google Scholar]
Pierdomenico M, Russo T, Ambroso S, Gori A, Martorelli E, D’Andrea L, Gili JM, Chiocci FL. 2018. Effects of trawling activity on the bamboo-coral Isidella elongata and the sea pen Funiculina quadrangularis along the Gioia Canyon (Western Mediterranean, southern Tyrrhenian Sea). Prog Oceanogr 169: 214–226. https://doi.org/10.1016/j.pocean.2018.02.019 [Google Scholar]
Purkey SG, Johnson GC. 2010. Warming of global abyssal and deep Southern Ocean waters between the 1990s and 2000s: Contributions to global heat and sea level rise budgets. J Clim 23: 6336–6351. https://doi.org/10.1175/2010JCLI3682.1 [Google Scholar]
Robert K, Jones DOB, Roberts JM, Huvenne VAI. 2016. Improving predictive mapping of deep-water habitats: Considering multiple model outputs and ensemble techniques. Deep Sea Res Part I Oceanogr Res Pap 113: 80–89. https://doi.org/10.1016/j.dsr.2016.04.008 [Google Scholar]
Rodríguez-Rodríguez D, Rodríguez J, Abdul Malak D, Nastasi A, Hernández P. 2016. Marine protected areas and fisheries restricted areas in the Mediterranean: assessing “actual” marine biodiversity protection coverage at multiple scales. Mar Policy 64: 24–30. https://doi.org/10.1016/j.marpol.2015.11.006 [Google Scholar]
Roger S, Vaz S, Tessier E, Cheret I, Verin Y, Ancellin J, Peres JM. 2024. Catch data from a 1949 trawl survey in Tunisia (South-West Mediterranean). SEANOE. https://doi.org/10.17882/99868 [Google Scholar]
Russo E, Monti MA, Mangano MC, Raffaetà A, Sarà G, Silvestri C, Pranovi F. 2020. Temporal and spatial patterns of trawl fishing activities in the Adriatic Sea (Central Mediterranean Sea, GSA17). Ocean Coast Manage 192: 105231. https://doi.org/10.1016/j.ocecoaman.2020.105231 [Google Scholar]
S, Vaz S, Tessier E, Cheret I, Verin Y, Ancellin J, Peres JM. 2024. Catch data from a 1949 trawl survey in Tunisia (South-West Mediterranean). SEANOE. https://doi.org/10.17882/99868 [Google Scholar]
Salgado-Hernanz PM, Regaudie-de-Gioux A, Antoine D, Basterretxea G. 2022. Pelagic primary production in the coastal Mediterranean Sea: variability, trends, and contribution to basin-scale budgets. Biogeosciences 19: 47–69. https://doi.org/10.5194/bg-19-47-2022 [Google Scholar]
Salomidi M, Gerovasileiou V, Stamouli C, Drakopoulou V, Otero MM, Jimenez C, Kiparissis S, Mytilineou Ch, Papadopoulou N, Smith CJ, Thasitis I, Anastasiadou Ch, Lefkaditou E, Makovsky Y, Schüler M. Deep-sea vulnerable benthic fauna, in: Otero M, Mytilineou C (Eds.), Deep-Sea Atlas of the Eastern Mediterranean Sea. IUCN-HCMR DeepEastMed Project, IUCN, Gland and Malaga, 2022, pp 123–144. [Google Scholar]
Salvalaggio V, Fabi G, Punzo E. 2016. Distribution of the sea pens Virgularia mirabilis and Funiculina quadrangularis (Cnidaria Anthozoa) in the northern and central Adriatic Sea. Rapport Du 41e Congres De La CIESM [Google Scholar]
Schickele A, Leroy B, Beaugrand G, Goberville E, Hattab T, Francour P, Raybaud V. 2020. Modelling European small pelagic fish distribution: Methodological insights. Ecol Model 416: 108902. https://doi.org/10.1016/j.ecolmodel.2019.108902 [Google Scholar]
Skliris N, Zika JD, Herold L, Josey SA, Marsh R. 2018. Mediterranean sea water budget long-term trend inferred from salinity observations. Clim Dyn 51: 2857–2876. https://doi.org/10.1007/s00382-017-4053-7 [Google Scholar]
Smith CJ, Gerovasileiou V, Mytilineou Ch, Jimenez C, Papadopoulou K, Salomidi M, Sakellariou D, Drakopoulou V, Otero M. 2022. Revisiting underwater surveys to uncover sites of conservation interest, in: Otero M, Mytilineou C (Eds.), Deep-Sea Atlas of the Eastern Mediterranean Sea. IUCN-HCMR DeepEastMed Project, IUCN, Gland and Malaga, pp. 146–171. [Google Scholar]
Smith KL, Ruhl HA, Kahru M, Huffard CL, Sherman AD. 2013. Deep ocean communities impacted by changing climate over 24 y in the abyssal northeast Pacific Ocean. Proc Natl Acad Sci USA 110: 19838–19841. https://doi.org/10.1073/pnas. 1315447110 [Google Scholar]
Sweetman AK, Thurber AR, Smith CR, Levin LA, Mora C, Wei CL, Gooday AJ, Jones DOB, Rex M, Yasuhara M, Ingels J, Ruhl HA, Frieder CA, Danovaro R, Würzberg L, Baco A, Grupe BM, Pasulka A, Meyer KS, Dunlop KM, Henry LA, Roberts JM. 2017. Major impacts of climate change on deep-sea benthic ecosystems. Elem Sci Anthropocene 5: 4. https://doi.org/10.1525/elementa.203 [Google Scholar]
Terribile K, Evans J, Knittweis L, Schembri PJ. 2016. Maximising MEDITS: Using data collected from trawl surveys to characterize the benthic and demersal assemblages of the circalittoral and deeper waters around the Maltese Islands (Central Mediterranean). Reg Stud Mar Sci 3:163–175. https://doi.org/10.1016/j.rsma.2015.07.006 [Google Scholar]
Thuiller W, Georges D, Engler R, Breiner F. 2023. biomod2: ensemble platform for species distribution modeling. R package ver 4.2-4 [Google Scholar]
Tudela S. 2004. Ecosystem effects of fishing in the Mediterranean: An analysis of the major threats of fishing gear and practices to biodiversity and marine habitats. Gen Fish Counc Medit Stud Rev 74. FAO, Rome, 44 pp. [Google Scholar]
UNGA. 2006. Resolution 61/105 adopted by the General Assembly on Sustainable fisheries, including through the 1995 Agreement for the Implementation of the Provisions of the United Nations Convention on the Law of the Sea of 10 December 1982 relating to the Conservation and Management of Straddling Fish Stocks and Highly Migratory Fish Stocks. UN Gen Assem Res 61/105. https://doi.org/10.18356/9789210021753 [Google Scholar]
Vasquez M, Ségeat B, Cordingley A, Tilby E, Wikström S, Ehrnsten E, Al Hamdani Z, Agnesi S, Andersen MS, Annunziatellis A, Askew N, Bekkby T, Bentes L, Daniels E, Doncheva V, Drakopoulou V, Ernstsen VB, Gonçalves J, Karvinen V, Laamanen-Nicolas L, Lillis H, Loukaidi V, Manca E, McGrath F, Mo G, Monteiro P, Muresan M, Nygard H, O’Keeffe E, Pelembe T, Radicioli M, Sakellariou D, Teaca A, Todorova V, Tunesi L, Woods H. 2023. EUSeaMap 2023, A European broad-scale seabed habitat map. https://doi.org/10.13155/97116 [Google Scholar]
Vaz S. 2018a. EPIBENGOL 2018 cruise, RV L’Europe. https://doi.org/10.17600/18000589 [Google Scholar]
Vaz S. 2018b. NOURMED. https://doi.org/10.18142/296 [Google Scholar]
Vaz S, Certain G, Hattab T, Jadaud A, Villeneuve R, Raphalen E, Metral L, Cheret I, Bourdeix JH, Tessier E. 2023. Suivi spatio-temporel des espèces démersales et benthiques: Rapport final de l’action 3 du Projet GOLDYS. https://doi.org/10.13155/96823 [Google Scholar]
Wang S, Murillo FJ, Kenchington E. 2022. Climate-change refugia for the bubblegum coral Paragorgia arborea in the Northwest Atlantic. Front Mar Sci 9: 863693. https://doi.org/10.3389/fmars.2022.863693 [Google Scholar]
Wright EP, Kemp K, Rogers AD, Yesson C. 2015. Genetic structure of the tall sea pen Funiculina quadrangularis in NW Scottish sea lochs. Mar Ecol 36: 659–667. https://doi.org/10.1111/maec.12174 [Google Scholar]

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