Marine Biodiversity

, Volume 49, Issue 2, pp 707–726 | Cite as

Bryozoan assemblages from two submarine caves in the Aegean Sea (Eastern Mediterranean)

  • Antonietta RossoEmail author
  • Vasilis Gerovasileiou
  • Rossana Sanfilippo
  • Adriano Guido
Original Paper


Although bryozoans are one of the dominant sessile phyla in Mediterranean marine caves, little information is available on the bryozoan diversity of this habitat in the eastern basin. In this study, bryozoan assemblages of two Aegean marine caves located at different depths and characterised by different morphology were studied. The examination of 30 quadrats (20 × 20 cm) scraped from the vertical walls and cave ceilings, at increasing distances from the entrances, yielded 74 taxa (67 living and 47 dead), half of which were shared by both caves. Cheilostomes largely prevailed (59 taxa) over cyclostomes and ctenostomes (14 and 1 species, respectively). Nodular to fungiform bioconstructions were formed by species with multilayered colonies in both caves. Differences in the taxonomic structure of bryozoan assemblages between the two caves as well as on the spatial variability of diversity, abundance and growth morphologies within each cave seem to be linked to the different cave morphology and associated environmental conditions. Moreover, a considerable number of taxa were reported for the first time from the Eastern Mediterranean and the marine cave habitat, supporting the idea that knowledge of their bryozoan diversity is far from being considered complete.


Bryozoa Biodiversity Bioconstructions Distribution Eastern Mediterranean Sea Holocene-to-Recent 



The authors would like to thank Eleni Voultsiadou and Drosos Koutsoubas for their support, Maria Sini for helping during fieldwork and Thanos Dailianis for providing underwater photographs. We would also like to thank Alfio Viola (University of Catania) for SEM assistance. This is contribution number 434 of the Catania Palaeontological Research Group.


This study was supported by the Research Funding Programme “Heracleitus II: Investing in knowledge society” (EU Social Fund and Greek national funds) and LifeWatchGreece research infrastructure (MIS 384676), which was funded by the Greek Government under the General Secretariat of Research and Technology (GSRT), ESFRI Projects, National Strategic Reference Framework (NSRF). VG also benefited from “Alexander S. Onassis Public Benefit Foundation” fellowship for postgraduate studies. Additional funds were also provided to AR by the University of Catania.

Compliance with ethical standards

Conflict of interest

The authors declare that they have no conflict of interest.

Ethical statements

All necessary permits for sampling and observational field studies have been obtained by the authors from the competent authorities and are mentioned in the acknowledgements, if applicable. This study has been approved by the Greek Ministry of Education, Lifelong Learning and Religious Affairs, through the acceptance of a Ph.D. project proposal. The locations studied are not privately owned nor protected in any way.


  1. Achilleos K, Jimenez C, Berning B, Petrou A (in press) Bryozoan diversity of Cyprus (Eastern Mediterranean): 4th century BCE until present with a particular focus on Monoporella bouchardii (Audouin & Savigny, 1826). In: Smidt R, Gordon DP and Reid C (eds) Proceedings of the XVII Conference of the International Bryozoology Association. Memoirs of the Association of Australasian PalaeontologistsGoogle Scholar
  2. Arko-Pijevac M, Benac Č, Kovačić M, Kirinčić M (2001) A submarine cave at the Island of Krk (North Adriatic Sea). Nat Croat 10(3):163–184Google Scholar
  3. Balduzzi A, Bianchi CN, Boero F, Cattaneo Vietti R, Pansini M, Sarà M (1989) The suspension-feeder communities of a Mediterranean sea cave. In: Ros JD (ed.), topics in marine biology. Sci Mar 53:387–395Google Scholar
  4. Balduzzi A, Rosso A (2003) Briozoi. In: Cicogna F, Bianchi NC, Ferrari G, Forti P (eds) Grotte marine: cinquant’anni di ricerca in Italia. Ministero dell’Ambiente e della Tutela del Territorio: Rapallo 195–202Google Scholar
  5. Berning B, Tilbrook KJ, Rosso A (2008) Revision of the north-eastern Atlantic and Mediterranean species of the genera Herentia and Therenia (Bryozoa: Cheilostomata). J Nat Hist 42(21–22):1509–1547. CrossRefGoogle Scholar
  6. Clarke KR, Gorley RN (2006) PRIMER v6: User Manual/Tutorial. PRIMER-E, PlymouthGoogle Scholar
  7. Di Geronimo I, Allegri L, Improta S, La Perna R, Rosso A, Sanfilippo R (1997) Spatial and temporal aspects of benthic thanatocoenoses in a Mediterranean infralittoral cave. Riv Ital Paleontol Stratigr 103:15–28Google Scholar
  8. Di Geronimo I, La Perna R, Rosso A, Sanfilippo R (1993) Popolamento e tanatocenosi bentonica della Grotta dell’Accademia (Ustica, Mar Tirreno meridionale). Nat Sicil 4(17):45–63Google Scholar
  9. Di Geronimo I, La Perna R, Rosso A, Sanfilippo R, Taddei-Ruggiero E (2000) Associazioni bentoniche da sedimenti di grotte carsiche in Sicilia. Speleol Iblea 8:97–102Google Scholar
  10. Fraschetti S, Boero F, Guarnieri G, Terlizzi A, Guidetti P, Bussotti S, Piraino S, De Vito D, Cormaci M, Furnari G, Catra M, Alongi G, Rosso A, Di Martino E, Ceccherelli G, Manconi R, Ledda F, Cattaneo-Vietti R, Cerrano C, Pantaleo U, Scinto A, Bavestrello G, Di Camillo CG, Betti F, Chemello R, Milazzo M, Graziano M, Di Franco A, Marchini A, Russo GF, Di Stefano F, Cimmino P (2010) Studio degli ambienti di grotte marine sommerse (Cod. 8330) nelle aree marine protette di Capo Caccia, Plemmirio e Isole Pelagie. Relazione finale. CoNISMa-Ministero dell’Ambiente e della Tutela del Territorio e del MareGoogle Scholar
  11. Gautier YV (1962) Recherches ecologiques sur les Bryozoaires Chilostomes en Méditerranée occidentale. Rec Trav Station Mar Endoume 38: 1–435Google Scholar
  12. Gerovasileiou V, Voultsiadou E (2015) Mediterranean marine caves as biodiversity reservoirs: a preliminary overview. In: Langar H, Bouafif C, Ouerghi A (eds) Proceedings of the 1st Mediterranean symposium on the conservation of dark habitats. RAC/SPA, Tunis, pp 45–50Google Scholar
  13. Gerovasileiou V, Rosso A (2016) Marine Bryozoa of Greece: an annotated checklist. Biodivers Data J 4:e10672CrossRefGoogle Scholar
  14. Gerovasileiou V, Voultsiadou E (2016) Sponge diversity gradients in marine caves of the eastern Mediterranean. J Mar Biol Assoc U K 96:407–416CrossRefGoogle Scholar
  15. Gerovasileiou V, Trygonis V, Sini M, Koutsoubas D, Voultsiadou E (2013) Three-dimensional mapping of marine caves using a handheld echosounder. Mar Ecol Prog Ser 486:13–22CrossRefGoogle Scholar
  16. Gerovasileiou V, Chintiroglou C, Vafidis D, Koutsoubas D, Sini M, Dailianis T, Issaris Y, Akritopoulou E, Dimarchopoulou D, Voultsiadou E (2015) Census of biodiversity in marine caves of the eastern Mediterranean Sea. Mediterr Mar Sci 16(1):245–265.
  17. Gerovasileiou V, Dimitriadis C, Arvanitidis C, Voultsiadou E (2017) Taxonomic and functional surrogates of sessile benthic diversity in Mediterranean marine caves. PLoS ONE 12(9):e0183707CrossRefGoogle Scholar
  18. Guido A, Heindel K, Birgel D, Rosso A, Mastandrea A, Sanfilippo R, Russo F, Peckman J (2013) Pendant bioconstructions cemented by microbial carbonate in submerged marine caves (Holocene, SE Sicily). Paleogeogr Palaeoclimatol Palaeoecol 388:166–180CrossRefGoogle Scholar
  19. Guido A, Jimenez C, Achilleos K, Rosso A, Sanfilippo R, Hadjioannou L, Russo F, Mastandrea A (2017) Geomicrobiology of the Kakoskali submarine cave (Cyprus, eastern Mediterranean). Facies 63:21. CrossRefGoogle Scholar
  20. Harmelin J-G (1969) Bryozoaires des grottes sous-marines obscures de la région Marseillaise: faunistique et écologie. Téthys 1:793–806Google Scholar
  21. Harmelin J-G (1976) Le sous-ordre des Tubuliporina (Bryozoaires Cyclostomes) en Méditerranée: écologie et systématique. Mem Inst océanogr Monaco 10:1–326Google Scholar
  22. Harmelin J-G (1980) Etablissement des communautés de substrats durs en milieu obscur. Résultats préliminaires d’une expérience à long terme en Méditerranée. Mem Biol Mar Ocean 10:29–52Google Scholar
  23. Harmelin J-G (1985) Bryozoan dominated assemblages in Mediterranean cryptic environments. In: Nielsen C, Larwood GP (eds) Bryozoan: Ordovician to recent. Olsen & Olsen, Fredensborg, pp 135–143Google Scholar
  24. Harmelin J-G (1986) Patterns in the distribution of bryozoans in the Mediterranean marine caves. Stygologia 2:10–25Google Scholar
  25. Harmelin J-G (1997) Diversity of bryozoans in a Mediterranean sublittoral cave with bathyal-like conditions: role of dispersal processes and local factors. Mar Ecol Prog Ser 153:139–152CrossRefGoogle Scholar
  26. Harmelin J-G (2000) Ecology of cave and cavity dwelling bryozoans. In: Herrera CA, Jackson JBC (eds) Proceedings of the 11th international Bryozoology association conference. Smithsonian Tropical Research Institute, Panama, pp 38–55Google Scholar
  27. Harmelin J-G (2003) Biodiversité des habitats cryptiques marins du parc national de Port-Cros (Méditerranée, France). Assemblages de bryozoaires d’une grotte sous-marine et des faces inférieures de pierres. Sci Rep Port-Cros Natl Park, France 19:101–115Google Scholar
  28. Harmelin J-G (2014a) Monoporella bouchardii (Audouin & Savigny, 1826) (Bryozoa, Cheilostomata): a forgotten taxon redescribed from the eastern Mediterranean material. Cah Biol Mar 55:91–99Google Scholar
  29. Harmelin J-G (2014b) Alien bryozoans in the eastern Mediterranean Sea–new records from the coasts of Lebanon. Zootaxa 3893(3):301–338CrossRefGoogle Scholar
  30. Harmelin J-G, Bitar G, Zibrowius H (2007) Schizoretepora hassi sp. nov. (Bryozoa: Phidoloporidae) from Lebanon and reappraisal of Scizotheca serratimargo (Hincks, 1886). Cah Biol Mar 48:179–186Google Scholar
  31. Harmelin J-G, Bitar G, Zibrowius H (2009) Smittinidae (Bryozoa, Cheilostomata) from coastal habitats of Lebanon (Mediterranean Sea), including new and non-indigenous species. Zoosystema 31(1):163–187CrossRefGoogle Scholar
  32. Harmelin J-G, Bitar G, Zibrowius H (2016) High xenodiversity versus low native biodiversity in the south-eastern Mediterranean: bryozoans from the coastal zone of Lebanon. Mediterr Mar Sci 17(2):417–439. CrossRefGoogle Scholar
  33. Harmelin J-G, Ostrovsky AN, Cáceres-Chamizo JP, Sanner J (2011) Bryodiversity in the tropics: taxonomy of Microporella species (Bryozoa Cheilostomata) with personate maternal zooids from Indian Ocean, Red Sea and southeast Mediterranean. Zootaxa 2798:1–30CrossRefGoogle Scholar
  34. Harmelin J-G, Vacelet J, Vasseur P (1985) Les grottes sous-marines obscures: un milieu extrême et un remarquable biotope refuge. Téthys 11:214–229Google Scholar
  35. Hayward PJ (1974) Studies on the cheilostome bryozoan fauna of the Aegean island of Chios. J Nat Hist 8:369–402Google Scholar
  36. Katsikatsos G, Migiros G, Triantaphyllis M, Mettos A (1986) Geological structure of internal Hellenides (E. Thessaly – SW Macedonia, Euboea – Attica – Northern Cyclades Islands and Lesbos), I.G.M.E, Geological and Geophysical Research, 191–212Google Scholar
  37. Kidwell SM, Bosence DW (1991) Taphonomy and time-averaging of marine shelly faunas. In: Allison PA, Briggs DEG (eds) Taphonomy: releasing the data locked in the fossil record. Plenum, New York, pp 115–209CrossRefGoogle Scholar
  38. Martí R, Uriz MJ, Ballesteros E, Turon X (2004) Benthic assemblages in two Mediterranean caves: species diversity and coverage as a function of abiotic parameters and geographic distance. J Mar Biol Assoc U K 84:557–572CrossRefGoogle Scholar
  39. Monteiro-Marques V (1981) Peuplements des planchers envasés de trois grottes sous-marines obscures de la région de Marseille. Etude préliminaire. Téthys 10:89–96Google Scholar
  40. Okamura B, Harmelin J-G, Jackson JBC (2001) Refuges revisited: enemies versus flow and feeding as determinants of sessile animal distribution and form. In: Jackson JBC, Lidgard S, McKinney FK (eds) Evolutionary patterns: growth, form, and tempo in the fossil record. University of Chicago Press, Chicago, pp 61–93Google Scholar
  41. Pérès JM, Picard J (1964) Nouveau manuel de bionomie benthique de la Mer Méditerranée. Rec Trav Station Mar Endoume 31:1–137Google Scholar
  42. Reverter Gil O, Souto J, Novosel M, Tilbrook K (2015) Adriatic species of Schizomavella (Bryozoa: Cheilostomata). J Nat Hist.
  43. Rosso A (1996) Popolamenti e tanatocenosi a Briozoi di fondi mobili circalitorali del Golfo di Noto (Sicilia SE). Nat Siciliano 20(3–4):189–225Google Scholar
  44. Rosso A, Di Martino E (2016) Mediterranean bryozoan biodiversity: an up-date. Mediterr Mar Sci 17(2):567–607. CrossRefGoogle Scholar
  45. Rosso A, Novosel M (2010) The genus Adeonella (Bryozoa, Ascophora) in the Mediterranean from Pliocene to recent, with description of two new species. J Nat Hist 44(27–28):1697–1727. CrossRefGoogle Scholar
  46. Rosso A, Di Martino E, Sanfilippo R, Di Martino V (2013a) Bryozoan communities and Thanatocoenoses from submarine caves in the Plemmirio marine protected area (SE Sicily). In: Ernst A, Schäfer P, Scholz J (eds) Bryozoan studies 2010, proceedings of the 15th IBA conference, Kiel, Germany. Lecture notes in earth system sciences, vol 143. Springer, Berlin, pp 251–269Google Scholar
  47. Rosso A, Sanfilippo R, Ruggieri R, Maniscalco R, Vertino A (2015) Exceptional record of submarine cave communities from the Pleistocene of Sicily (Italy). Lethaia 48:133–144. CrossRefGoogle Scholar
  48. Rosso A, Sanfilippo R, Sciuto F (2014) Open shelf soft bottom bryozoan communities from the Ciclopi marine protected area (E Sicily, Mediterranean). In: Rosso A, Wyse Jackson PN, Porter J (eds). Bryozoan studies 2013. Proceedings of the 16th Conference of the International Bryozoological Association, Catania: Studi Trentini di Scienze Naturali, 94: 195–207Google Scholar
  49. Rosso A, Sanfilippo R, Taddei-Ruggiero E, Di Martino E (2013b) Serpuloidean, bryozoan and brachiopod faunas from submarine caves in Sicily. Boll Soc Paleontol Ital 52(3):167–176Google Scholar
  50. Sanfilippo R, Rosso A, Guido A, Gerovasileiou V (2017) Serpulid communities from two marine caves in the Aegean Sea, eastern Mediterranean. J Mar Biol Assoc U K.
  51. Sanfilippo R, Rosso A, Guido A, Mastandrea A, Russo F, Ryding R, Taddei Ruggero E (2015) Metazooan/microbial biostalactites from modern submarine caves in the Mediterranean Sea. Mar Ecol 36:1277–1293CrossRefGoogle Scholar
  52. Taddei Ruggiero E, Annunziata G, Rosso A, Sanfilippo R (1996) Il benthos della Grotta sottomarina dell’Isca (Penisola Sorrentina): evidenze faunistiche della sua evoluzione recente. VII Congresso Nazionale della Società Italiana di Ecologia 17:329–332Google Scholar
  53. Zabala M, Gili JM (1985) Distribution des Bryozoaires le long d’un gradient sédimentaire dans deux grottes sous-marines du littoral de Majorque. Commission Internationale pour L’exploration Scientifique de la Mer Méditerranée 29:137–140Google Scholar
  54. Zabala M, Rieira T, Gili JM, Barange M, Lobo A, Peñuelas J (1989) Water flow, Trophic depletion, and benthic macrofauna impoverishment in a submarine cave from the western Mediterranean. PSZNI Mar Ecol 10:271–287CrossRefGoogle Scholar

Copyright information

© Senckenberg Gesellschaft für Naturforschung and Springer-Verlag GmbH Germany, part of Springer Nature 2018

Authors and Affiliations

  1. 1.Dipartimento di Scienze Biologiche, Geologiche e Ambientali, Corso ItaliaUniversità di CataniaCataniaItaly
  2. 2.CoNISMa (Consorzio Nazionale Interuniversitario per le Scienze del Mare), URL CataniaCataniaItaly
  3. 3.Institute of Marine Biology, Biotechnology and AquacultureHellenic Centre for Marine ResearchHeraklionGreece
  4. 4.Dipartimento di BiologiaEcologia e Scienze della TerraCosenzaItaly

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