Taxonomic and morphological descriptors reveal high benthic temporal variability in a Mediterranean marine submerged cave over a decade


The effectiveness of monitoring programs in the marine environment is often limited by the scarcity of information concerning medium/long-term time series observations (years-decades) in complex and variable systems. Marine submerged caves, in particular, have received relatively low research effort, as they are confined and difficult to access ecosystems. In this study, we describe the spatio-temporal variations of sessile benthic assemblages along the exterior-interior axis of the Cerro-Gordo submarine cave (Granada, Spain), from 2007 to 2016 using taxonomic (e.g., major taxonomic groups) and morphological (i.e., growth forms) descriptors. In contrast to our initial expectations (higher stability in the inner cave sector), significant temporal variations of sessile community structure and morphology occurred in both external and internal cave sectors. In addition, higher spatial differences were encountered among opposite cave walls in the inner dark sector, which is probably explained by small-scale variability in topography and siltation rate. In the outer semi-dark sector, community structure was more heterogeneous, although generally similar between opposite walls. Among the descriptors used, the morphological one was able to detect greater spatio-temporal variation in the outer semi-dark sector, while the taxonomic descriptor was more suitable in the inner dark sector. These results could provide a basis for future monitoring programs.

This is a preview of subscription content, log in to check access.

Access options

Buy single article

Instant unlimited access to the full article PDF.

US$ 39.95

Price includes VAT for USA

Subscribe to journal

Immediate online access to all issues from 2019. Subscription will auto renew annually.

US$ 199

This is the net price. Taxes to be calculated in checkout.

Fig. 1
Fig. 2
Fig. 3
Fig. 4
Fig. 5


  1. Abdulla, A., M. Gomei, É. Maison & C. Piante, 2008. Status of Marine Protected Areas in the Mediterranean Sea. IUCN, Málaga, and WWF, Paris.

  2. Anderson, M. J., 2001. A new method for non-parametric multivariate analysis of variance. Austral Ecology 26: 32–46.

  3. Anderson, M. J., C. E. Diebel, W. M. Blom & T. J. Landers, 2005. Consistency and variation in kelp holdfast assemblages: spatial patterns of biodiversity for the major phyla at different taxonomic resolutions. Journal of Experimental Marine Biology and Ecology 320: 35–56.

  4. Anderson, M. J., R. N. Gorley & K. R. Clarke, 2008. PERMANOVA + for PRIMER. Guide to software and statistical methods. PRIMER-E Ltd, Plymouth.

  5. Baldacconi, R. & E. Trainito, 2013. Spugne del Mediterraneo. Edizioni il Castello.

  6. Benedetti-Cecchi, L., L. Airoldi, M. Abbiati & F. Cinelli, 1998. Spatial variability in the distribution of sponges and cnidarians in a sublittoral marine cave with sulphur-water springs. Journal of the Marine Biological Association of the United Kingdom 78: 43–58.

  7. Bianchi, C. N. & C. Morri, 1994. Studio bionomico comparativo di alcune grotte marine sommerse: definizione di una scala di confinamento. Istituto Italiano di Speleologia Memoria 6: 107–123.

  8. Bianchi, C. N., R. Cattaneo-Vietti, F. Cinelli, C. Morri & M. Pansini, 1996. Lo studio biologico delle grotte sottomarine del Mediterraneo: conoscenze attuali e prospettive. Bollettino dei Musei e degli Istituti Biologici dell’Università di Genova 60(61): 41–69.

  9. Bibiloni, M. A., M. J. Uriz & J. M. Gili, 1989. Sponge communities in three submarine caves of the Balearic Islands (Western Mediterranean): adaptations and faunistic composition. Marine Ecology 10: 317–334.

  10. Boury-Esnault, N., J. G. Harmelin & J. Vacelet, 1993. Les abysses méditerranéennes à vingt mètres de profondeur. La recherche 24: 848–851.

  11. Bussotti, S., A. Terlizzi, S. Fraschetti, G. Belmonte & F. Boero, 2006. Spatial and temporal variability of sessile benthos in shallow Mediterranean marine caves. Marine Ecology Progress Series 325: 109–119.

  12. Carballo, J. L. & J. C. Garcia-Gómez, 1994. Esponjas del Estrecho de Gibraltar y áreas próximas, con nuevas aportaciones para la fauna Iberica [Porifera from the Straits of Gibraltar and nearby areas, new species of the Iberian fauna]. Cahiers de Biologie Marine 35: 193–211.

  13. Carballo, J. L., S. Naranjo & J. C. García-Gómez, 1997. Where does the Mediterranean Sea begin? Zoogeographical affinities of the littoral sponges of the Straits of Gibraltar. Journal of Biogeography 24: 223–232.

  14. Cebrián, P. E. & S. E. Ballesteros, 2003. Zonation patterns of benthic communities in an upwelling area from de western Mediterranean (La Herradura, Alboran Sea). Scientia Marina 68: 69–84.

  15. Chevaldonné, P. & C. Lejeusne, 2003. Regional warming-induced species shift in north-west Mediterranean marine caves. Ecology Letters 6: 371–379.

  16. Cinelli, F., E. Fresi, L. Mazzella, M. Pansini, R. Pronzato & A. Svoboda, 1977. Distribution of benthic phyto-and zoocoenoses along a light gradient in a superficial marine cave. Biology of benthic organisms. Pergamon Press, Oxford: 173–183.

  17. Clarke, K. R. & R. N. Gorley, 2006. PRIMER v6: User Manual/Tutorial. PRIMER-E Ltd, Plymouth.

  18. Coll, M., C. Piroddi, J. Steenbeek, K. Kaschner, F. B. R. Lasram, J. Aguzzi, E. Ballesteros, C. N. Bianchi, J. Corbera, T. Dailianis, R. Danovaro, M. Estrada, C. Froglia, B. S. Galil, J. M. Gasol, R. Gertwagen, J. Gil, F. Guilhaumon, K. Kesner-Reyes, M. S. Kitsos, A. Koukouras, N. Lampadariou, E. Laxamana, C. M. López-Fé de la Cuadra, H. K. Lotze, D. Martin, D. Mouillot, D. Oro, S. Raicevich, J. Rius-Barile, J. I. Saiz-Salinas, C. San Vicente, S. Somot, J. Templado, X. Turon, D. Vafidis, R. Villanueva & E. Voultsiadou, 2010. The biodiversity of the Mediterranean Sea: estimates, patterns, and threats. PLoS ONE 5: e11842.

  19. Costa, G., F. Betti, E. Nepote, R. Cattaneo-Vietti, M. Pansini, G. Bavestrello & M. Bertolino, 2018. Sponge community variations within two semi-submerged caves of the Ligurian Sea (Mediterranean Sea) over a half-century time span. The European Zoological Journal 85: 382–392.

  20. De Goeij, J. M., D. Van Oevelen, M. J. Vermeij, R. Osinga, J. J. Middelburg, A. F. de Goeij & W. Admiraal, 2013. Surviving in a marine desert: the sponge loop retains resources within coral reefs. Science 342: 108–110.

  21. Delgado, M., 1990. Phytoplankton distribution along the Spanish coast of the Alboran Sea. Scientia Marina 54: 169–178.

  22. Di Franco, A., G. Ferruzza, P. Baiata, R. Chemello & M. Milazzo, 2010. Can recreational scuba divers alter natural gross sedimentation rate? A case study from a Mediterranean deep cave. ICES Journal of Marine Science 67: 871–874.

  23. Dimarchopoulou, D., V. Gerovasileiou & E. Voultsiadou, 2018. Spatial variability of sessile benthos in a semi-submerged marine cave of a remote Aegean Island (eastern Mediterranean Sea). Regional Studies in Marine Science 17: 102–111.

  24. Directive, 1992. Council Directive 92/43/EEC of 21 May 1992 on the conservation of natural habitats and of wild fauna and flora. Official Journal of the European Communities.

  25. Directive, 2000. Directive 2000/60/EC of the European Parliament and of the Council of 23 October 2000 establishing a framework for Community action in the field of water policy. Official Journal of the European Communities.

  26. Garrabou, J., E. Sala, C. Linares, J. B. Ledoux, I. Montero-Serra, J. M. Dominici, S. Kipson, N. Teixidó, E. Cebrian, D. K. Kersting & J. G. Harmelin, 2017. Re-shifting the ecological baseline for the overexploited Mediterranean red coral. Scientific Reports 7: 42404.

  27. Gerovasileiou, V. & E. Voultsiadou, 2014. Mediterranean marine caves as biodiversity reservoirs: a preliminary overview. In Bouafif, C., Langar H. & A. Ouerghi (eds), Proceedings of the 1st Mediterranean Symposium on the Conservation of Dark Habitats. RAC/SPA, Tunis: 45–50.

  28. Gerovasileiou, V., A. Martínez, F. Álvarez, G. Boxshall, W. F. Humphreys, D. Jaume, L. E. Becking, G. Muricy, P. J. van Hengstum, H. Yamasaki, N. Bailly & T. M. Iliffe, 2019. World Register of Marine Cave Species (WoRCS). [Last Accessed on February 2019].

  29. Gerovasileiou, V. & E. Voultsiadou, 2012. Marine Caves of the Mediterranean Sea: a Sponge Biodiversity Reservoir within a Biodiversity Hotspot. PLoS ONE 7: e39873.

  30. Gerovasileiou, V. & E. Voultsiadou, 2016. Sponge diversity gradients in marine caves of the Eastern Mediterranean. Journal of the Marine Biological Association of the United Kingdom 96: 407–416.

  31. Gerovasileiou, V., V. Trygonis, M. Sini, D. Koutsoubas & E. Voultsiadou, 2013. Three-dimensional mapping of marine caves using a handheld echosounder. Marine Ecology Progress Series 486: 13–22.

  32. Gerovasileiou, V., A. Martínez, F. Álvarez, G. Boxshall, W. Humphreys, D. Jaume, L. Becking, G. Muricy, P. van Hengstum, S. Dekeyzer, W. Decock, B. Vanhoorne, L. Vandepitte, N. Bailly & T. Iliffe, 2016. World Register of marine Cave Species (WoRCS): a new Thematic Species Database for marine and anchialine cave biodiversity. Research Ideas and Outcomes 2: e10451.

  33. Gerovasileiou, V., C. Dimitriadis, C. Arvanitidis & E. Voultsiadou, 2017. Taxonomic and functional surrogates of sessile benthic diversity in Mediterranean marine caves. PLoS ONE 12: e0183707.

  34. Gerovasileiou, V., T. Dailianis, M. Sini, M. Otero, C. Numa, S. Katsanevakis & E. Voultsiadou, 2018. Assessing the regional conservation status of sponges (Porifera): the case of the Aegean ecoregion. Mediterranean Marine Science 19: 526–537.

  35. Giakoumi, S., M. Sini, V. Gerovasileiou, T. Mazor, J. Beher, H. P. Possingham, A. Abdulla, M. E. Çinar, P. Dendrinos, A. C. Gucu, A. A. Karamanlidis, P. Rodic, P. Panayotidis, E. Taskin, A. Jaklin, E. Voultsiadou, C. Webster, A. Zenetos & S. Katsanevakis, 2013. Ecoregion-based conservation planning in the Mediterranean: dealing with large-scale heterogeneity. PLoS ONE 8: e76449.

  36. Gofas, S., C. Salas & D. Moreno, 2011. Moluscos marinos de Andalucía. Universidad de Málaga, Servicio de Publicaciones e Intercambio Científico, Málaga.

  37. Grenier, M., C. Ruiz, M. Fourt, M. Santonja, M. Dubois, M. Klautau, J. Vacelet, N. Boury-Esnault & T. Pérez, 2018. Sponge inventory of the French Mediterranean waters, with an emphasis on cave-dwelling species. Zootaxa 4466: 205–228.

  38. Guarnieri, G., A. Terlizzi, S. Bevilacqua & S. Fraschetti, 2012. Increasing heterogeneity of sensitive assemblages as a consequence of human impact in submarine caves. Marine Biology 159: 1155–1164.

  39. Guido, A., K. Heindel, D. Birgel, A. Rosso, A. Mastandrea, R. Sanfilippo & J. Peckmann, 2013. Pendant bioconstructions cemented by microbial carbonate in submerged marine caves (Holocene, SE Sicily). Palaeogeography, Palaeoclimatology, Palaeoecology 388: 166–180.

  40. Harmelin, J. G., 1969. Bryozoaires des grottes sous-marines obscures de la région marseillaise, faunistique et écologie. Téthys 1: 793–806.

  41. Harmelin, J. G., 1980. Etablissement des communautes de substrats durs en milieu obscur. Resultats preliminaires d’une experience à long terme en Mediterranée. Memorie di Biologia Marina e di Oceanografia 10: 29–52.

  42. Harmelin, J. G., 1986. Patterns in the distribution of bryozoans in the Mediterranean marine caves. Stygologia 2: 10–25.

  43. Harmelin, J. G., J. Vacelet & P. Vasseur, 1985. Les grottes sous-marines obscures: un milieu extreme et un remàrquable biotope refuge. Téthys 11: 214–229.

  44. Hofrichter, R. (coord.), 2005. El mar Mediterráneo. Fauna, flora, ecología, Vol II/1: Guía sistemática y de identificación. Ediciones Omega, Sabadell.

  45. IBERFAUNA, 2018. El Banco de Datos de la Fauna Ibérica. Museo Nacional de Ciencias Naturales (CSIC). [Last Accessed on November 2018].

  46. Iliffe, T. M., D. J. Timothy & S. B. Martin, 1984. Organic pollution of an inland marine cave from Bermuda. Marine Environmental Research 12: 173–189.

  47. Lafargue, F., A. A. Ramos, X. Turon, B. Banaigs & M. Wahl, 1986. The littoral ascidians of the Spanish mediterranean. I. From Port Bou to the Islas Medas. Vie et Milieu 36: 133–139.

  48. Lecher, A. L. & K. R. M. Mackey, 2018. Synthesizing the effects of submarine groundwater discharge on Marine Biota. Hydrology 5(4): 60.

  49. Logan, A., A. Tomašových, M. Zuschin & B. Grill, 2008. Recent brachiopods from the Red Sea and Gulf of Aden. Fossils and Strata 54: 299–309.

  50. Magurran, A. E., S. R. Baillie, S. T. Buckland, J. M. Dick, D. A. Elston, E. M. Scott, R. I. Smith, P. J. Somerfield & A. D. Watt, 2010. Long-term datasets in biodiversity research and monitoring: assessing change in ecological communities through time. Trends in Ecology and Evolution 25: 574–582.

  51. Maldonado, M., L. Sánchez-Tocino & C. Navarro, 2010. Recurrent disease outbreaks in corneous demosponges of the genus Ircinia: epidemic incidence and defense mechanisms. Marine Biology 157: 1577–1590.

  52. Mancenido, M. O. & R. Gourvennec, 2007. A reappraisal of feeding current systems inferred for spire-bearing brachiopods. Earth and Environmental Science Transactions of the Royal Society of Edinburgh 98: 345–356.

  53. MAPAMA, 2018. Estado de las masas de agua subterránea PHC 2009-2015. [online] Available at: Dirección General del Agua (DGA), Ministerio de Agricultura y Pesca, Alimentación y Medio Ambiente (MAPAMA). Gobierno de España [Last Accessed on November 2018].

  54. Martí, R., M. J. Uriz, E. Ballesteros & X. Turon, 2004. Benthic assemblages in two Mediterranean caves: species diversity and coverage as a function of abiotic parameters and geographic distance. Journal of the Marine Biological Association of the United Kingdom 84: 557–572.

  55. Montefalcone, M., G. De Falco, E. Nepote, M. Canessa, M. Bertolino, G. Bavestrello, C. Morri & C. N. Bianchi, 2018. Thirty year ecosystem trajectories in a submerged marine cave under changing pressure regime. Marine Environmental Research 137: 98–110.

  56. Navarro-Barranco, C., J. M. Guerra-García, L. Sánchez-Tocino & J. C. García-Gómez, 2012. Soft-bottom crustacean assemblages in Mediterranean marine caves: the cave of Cerro Gordo (Granada, Spain) as case study. Helgoland Marine Research 66: 567–576.

  57. Navarro-Barranco, C., J. M. Guerra-García, L. Sánchez-Tocino, P. Jiménez-Prada, S. Cea & J. C. García-Gómez, 2013. Soft-bottom diversity patterns in marine caves; Lessons from crustacean community. Journal of Experimental Marine Biology and Ecology 446: 22–28.

  58. Navarro-Barranco, C., J. M. Guerra-García, L. Sánchez-Tocino, M. Ros, M. Florido & J. C. García-Gómez, 2015. Colonization and successional patterns of the mobile epifaunal community along an environmental gradient in a marine cave. Marine Ecology Progress Series 521: 105–115.

  59. Nepote, E., C. N. Bianchi, C. Morri, M. Ferrari & M. Montefalcone, 2017. Impact of a harbour construction on the benthic community of two shallow marine caves. Marine Pollution Bulletin 114: 35–45.

  60. Ocaña, A., L. Sánchez Tocino & P. J. López-González, 2000. Consideraciones faunísticas y biogeográficas de los antozoos (Cnidaria: Anthozoa) de la costa de Granada (Mar de Alborán). Zoologica baetica 11: 51–65.

  61. Okamura, B., J. G. Harmelin & J. B. C. Jackson, 2001. Refuges revisited: enemies versus flow and feeding as determinants of sessile animal distribution and form. In Jackson, J. B. C., S. Lidgard & F. K. Mckinney (eds), Evolutionary Patterns: Growth, Form and Tempo in the Fossil Record. University of Chicago Press, Chicago: 61–93.

  62. Pansini, M., 1987. Littoral demosponges from the banks of the Strait of Sicily and the Alboran Sea. In Vacelet, J. & N. Boury-Esnault (eds), Taxonomy of Porifera from the N.E. Atlantic and the Mediterranean Sea. NATO Advanced Science Institutes Series G, Ecological Sciences. Springer, Heidelberg: 149–185.

  63. Parravicini, V., C. Morri, G. Ciribilli, M. Montefalcone, G. Albertelli & C. N. Bianchi, 2009. Size matters more than method: visual quadrats versus photography in measuring human impact on Mediterranean rocky reef communities. Estuarine, Coastal and Shelf Science 81: 359–367.

  64. Parravicini, V., P. Guidetti, C. Morri, M. Montefalcone, M. Donato & C. N. Bianchi, 2010. Consequences of sea water temperature anomalies on a Mediterranean submarine cave ecosystem. Estuarine, Coastal and Shelf Science 86: 276–282.

  65. Pohlman, J. W., T. M. Iliffe & L. A. Cifuentes, 1997. A stable isotope study of organic cycling and the ecology of an anchialine cave ecosystem. Marine Ecology Progress Series 155: 17–27.

  66. Puertos del Estado, 2018. Prediccion de oleaje, nivel del mar; Boyas y mareografos. Datos históricos, SIMAR: 2042080 and SIMAR: Málaga [online]. Ministerio de Fomento, Gobierno de España. [Last Accessed on November 2018].

  67. Quinn, J. F., 1982. Competitive hierarchies in marine benthic communities. Oecologia 54:129–135

  68. Radolović, M., T. Bakran-Petricioli, D. Petricioli, M. Surić & D. Perica, 2015. Biological response to geochemical and hydrological processes in a shallow submarine cave. Mediterranean Marine Science 16: 305–324.

  69. Rastorgueff, P. A., D. Bellan-Santini, C. N. Bianchi, S. Bussotti, P. Chevaldonné, P. Guidetti & S. Ruitton, 2015. An ecosystem-based approach to evaluate the ecological quality of Mediterranean undersea caves. Ecological Indicators 54: 137–152.

  70. Riisgård, H. U. & P. S. Larsen, 2000. Comparative ecophysiology of active zoobenthic filter feeding, essence of current knowledge. Journal of Sea Research 44: 169–193.

  71. Rodríguez-Prieto, C., E. Ballesteros, F. Boisset & J. Afonso-Carrillo, 2013. Guía de las macroalgas y fanerógamas marinas del Mediterráneo occidental. Omega, Barcelona.

  72. Rosso, A. & E. Di Martino, 2016. Bryozoan diversity in the Mediterranean Sea: an update. Mediterranean Marine Science 17: 567–607.

  73. Rosso A., Di Martino E., Sanfilippo R. & V. Di Martino, 2013a. Bryozoan Communities and Thanatocoenoses from Submarine Caves in the Plemmirio Marine Protected Area (SE Sicily). In Ernst A., P. Schäfer, & J. Scholz (eds), Bryozoan Studies 2010. Proceedings of the 15th IBA Conference, 2010 Kiel, Germany. Lecture Notes in Earth System Sciences, Vol 143. Springer, Berlin: 251–269.

  74. Rosso, A., R. Sanfilippo, Ruggiero E. Taddei & E. Di Martino, 2013b. Faunas and ecological groups of Serpuloidea, Bryozoa and Brachiopoda from submarine caves in Sicily (Mediterranean Sea). Bollettino Società Paleontologica Italiana 52: 167–176.

  75. Rosso, A., R. Sanfilippo & F. Matrototaro, 2015. Bryozoan and serpulid distribution pattern on deep/water slab (Bari Canyon, Adriatic Sea). In Bouafif, C., Langar, H. & A. Ouerghi (eds), Proceedings of the 1st Mediterranean Symposium on the conservation of Dark Habitats. RAC/SPA, Tunis: 79–80.

  76. Rosso, A., V. Gerovasileiou, R. Sanfilippo & A. Guido, 2019. Bryozoan assemblages from two submarine caves in the Aegean Sea (Eastern Mediterranean). Marine Biodiversity 49: 707–726.

  77. Sala, E., J. Garrabou & M. Zabala, 1996. Effects of diver frequentation on Mediterranean sublittoral populations of the bryozoan Pentapora fascialis. Marine Biology 126: 451–459.

  78. Sánchez-Tocino, L., 2018. El litoral de Granada, Universidad de Granada (UGR), Spain. [Last Accessed on November 2018].

  79. Sánchez-Tocino, L. & J. M. Tierno de Figueroa, 2016. Mass mortality event of the sponges Ircinia variabilis (Schmidt, 1862) and Sarcotragus fasciculatus (Pallas, 1766) (Porifera, Demospongia) in the southern Spanish coast. Zoologica Baetica 27: 39–46.

  80. Sanfilippo, R., A. Rosso, A. Guido, A. Mastandrea, F. Russo, R. Riding & E. R. Taddei, 2015. Metazoan/microbial biostalactites from present-day submarine caves in the Mediterranean Sea. Marine Ecology 36: 1277–1293.

  81. Sanfilippo, R., A. Rosso, A. Guido & V. Gerovasileiou, 2017. Serpulid communities from two marine caves in the Aegean Sea, eastern Mediterranean. Journal of the Marine Biological Association of the United Kingdom 97: 1059–1068.

  82. Sarà, M., 1976. Il popolamento delle grotte marine: interesse di una salvaguardia. Pubblicazioni della Stazione Zoologica di Napoli 40: 502–505.

  83. Schneider, D. C., 1994. Quantitative Ecology: Spatial and Temporal Scaling. Academic Press, San Diego.

  84. Sousa, W. P., 1984. The role of disturbance in natural communities. Annual review of ecology and systematics 15: 353–391

  85. SPA/RAC–UN Environment/MAP & OCEANA, 2017. Guidelines for inventorying and monitoring of dark habitats in the Mediterranean Sea. SPA/RAC—Deep Sea Lebanon Project, Tunis.

  86. Templado, J., 1993. Fauna marina circalitoral del sur de la Península Ibérica: resultados de la campaña oceanográfica "Fauna I". Editorial CSIC-CSIC Press.

  87. Terrón-Sigler, A., Moreno-Tempestini, L., Jiménez-Martínez, B., Aguilar-Domínguez, M. D., Gutiérrez-Alba, V. & D. León-Muéz, 2008. El coral naranja (Astroides calycularis); estudio, dinámica de población y medidas de gestión de un recurso natural endémico. In: XV Simposio Ibérico de Estudios de Biología Marina, Funchal.

  88. Terrón-Sigler, A., D. León-Muez, P. Penalver-Duque & F. Espinosa, 2016. The effects of SCUBA diving on the endemic Mediterranean coral Astroides calycularis. Ocean & Coastal Management 122: 1–8.

  89. Trygonis, V. & M. Sini, 2012. photoQuad: a dedicated seabed image processing software, and a comparative error analysis of four photoquadrat methods. Journal of Experimental Marine Biology and Ecology 424: 99–108.

  90. Turnhout, E., M. Hisschemoller & H. Eijsackers, 2007. Ecological indicators: between the two fires of science and policy. Ecological Indicators 7: 215–228.

  91. UNEP-MAP-RAC, SPA, 2014. SPAMI list. UNEP-MAP-RAC/SPA, Tunis.

  92. UNEP-MAP-RAC/SPA, 2015. Action Plan for the Conservation of Habitat and Species Associated with Seamounts, Underwater Caves and Canyons, Aphotic Hard Beds and Chemo-synthetic Phenomena in the Mediterranean Sea. Dark Habitats Action Plan. RAC/SPA, Tunis.

  93. Zabala, M., T. Riera, J. M. Gili, M. Barange, A. Lobo & J. Peñuelas, 1989. Water flow, trophic depletion, and benthic macrofauna impoverishment in a submarine cave from the Western Mediterranean. Marine Ecology 10: 271–287.

Download references


We would like to thank Enrique Quintero-Torres (Venezuelan Institute for Scientific Research) and César Megina (University of Seville) for their help in data analysis. We are also grateful to Carlos González, Raúl Rodríguez Roig and “Open Water La Herradura” diving center for their collaboration during field samplings. The authors would also like to thank two anonymous reviewers and the Associate Editor-in-chief Diego Fontaneto for their constructive comments.

Author information

Correspondence to Juan Sempere-Valverde.

Additional information

Publisher's Note

Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

Handling editor: Diego Fontaneto

Electronic supplementary material

Below is the link to the electronic supplementary material.

Supplementary material 1 (MP4 68800 kb)

Supplementary material 1 (MP4 68800 kb)

Rights and permissions

Reprints and Permissions

About this article

Verify currency and authenticity via CrossMark

Cite this article

Sempere-Valverde, J., Lorenzo, Á.S., Espinosa, F. et al. Taxonomic and morphological descriptors reveal high benthic temporal variability in a Mediterranean marine submerged cave over a decade. Hydrobiologia 839, 177–194 (2019).

Download citation


  • Submerged caves
  • Benthic community
  • Morphological descriptors
  • Taxonomic descriptors
  • Growth forms