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A Preliminary Survey of Marine A preliminary survey of marine cave habitats in the Maltese Islands L Knittweis, P Chevaldonné, Alexander Ereskovsky, J Schembri, J.A. Borg To cite this version: L Knittweis, P Chevaldonné, Alexander Ereskovsky, J Schembri, J.A. Borg. A preliminary sur- vey of marine cave habitats in the Maltese Islands. Xjenza Online, 2015, 3 (2), pp.153-164. 10.7423/XJENZA.2015.2.07. hal-01463733 HAL Id: hal-01463733 https://hal.archives-ouvertes.fr/hal-01463733 Submitted on 13 Apr 2018 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Knittweis, L., Chevaldonn´e,P., Ereskovsky, A., Schembri, P. J., and Borg, J. A. (2015).Xjenza Online, 3:153{164. Xjenza Online - Journal of The Malta Chamber of Scientists www.xjenza.org DOI: 10.7423/XJENZA.2015.2.07 Research Article A preliminary survey of marine cave habitats in the Maltese Islands L. Knittweis∗1, P. Chevaldonn´e2, A. Ereskovsky2;3, P. J. Schembri1, J. A. Borg1 1Department of Biology, Faculty of Science, University of Malta, Msida MSD 2080, Malta 2Institut M´editerran´eende Biodiversit´eet d'Ecologie Marine et Continentale (IMBE), CNRS, IRD, Aix Marseille Universit´e,Avignon Universit´e,Station Marine d'Endoume, 13007 Marseille, France 3Faculty of Biology, Saint-Petersburg State University, 7/9 Universitetskaya Emb. St. Petersburg 199034, Russia Abstract. The Mediterranean Sea is a hotspot for mar- 1 Introduction ine biodiversity. Past studies of Mediterranean marine The Mediterranean Sea is a hotspot for marine biod- caves have revealed the unique biocoenotic and ecolo- iversity and ranks among the most important of such gical characteristics of these habitats, which are protec- regions worldwide (Bianchi et al., 2012), although much ted by European Union legislation. The Maltese Islands research remains to be done to assess and monitor the have an abundance of partially and fully submerged biodiversity it supports, and to better understand its marine caves with different geomorphological character- marine ecosystems. Despite numerous national and in- istics, yet there have been no systematic studies on these ternational programs of research on the flora and fauna habitats and their associated species. This study is a of the Mediterranean Sea, its biodiversity still remains first synthesis of existing information on the biotic as- insufficiently studied (Coll et al., 2010). The description semblages and physical characteristics of Maltese mar- of new species, especially of invertebrates from poorly ine caves. The work combines a review of the available known groups or from groups which thrive in habitats information with a preliminary survey of some marine that are difficult to explore such as the deep sea and caves in Gozo, during which several species were recor- marine caves, is an ongoing process and new discoveries ded for the first time for the Maltese Islands. Character- continually add to previous estimates of overall Mediter- istic species recorded from local marine caves are high- ranean species diversity. In particular, increased effort lighted, including several species of red and brown algae, to study the species diversity and the ecology of poorly sessile invertebrates including bryozoans, ascidians and known habitats is required, as are long-term monitoring sponges, and mobile forms including crustaceans and programs of marine species and habitats. fish. A marked zonation from the cave entrance to the Past studies of Mediterranean marine caves have re- inside of the caves was identified: photophilic algae at vealed the unique biocoenotic and ecological charac- the mouth of the cave are progressively replaced by more teristics of these habitats. Sessile benthic species as- sciaphilic species, followed by a middle section domin- sociated with cave habitats typically show a marked ated by sessile invertebrates, and then a completely dark zonation from the cave entrance to the inward end of inner section that is mostly devoid of sessile organisms. the cave: photophilic algae at the mouth of the cave Several species protected by national and international are rapidly replaced by encrusting coralline algae and legislation were found to occur. other more sciaphilic species. Thereafter a middle sec- Keywords: Mediterranean marine caves, sciaphilic, tion tends to be dominated by sessile invertebrates such cave biota, sessile biotic assemblages, species distribu- as sponges, corals, and bryozoans, while the completely tion patterns dark inner section tends to be mostly devoid of sessile organisms (Laborel & Vacelet, 1958; Riedl, 1966). This characteristic zonation is thought to be a consequence of the strong environmental gradients that exist, especially those related to light intensity and water movement, as one proceeds inwards from the cave mouth (Bussotti, *Correspondence to: L. Knittweis ([email protected]) c 2015 Xjenza Online A preliminary survey of marine cave habitats in the Maltese Islands 154 Terlizzi, Fraschetti, Belmonte & Boero, 2006; Gili, Ri- and eastern basins of the Mediterranean. From the era & Zabala, 1986; Harmelin, Vacelet & Vasseur, 1985; point of view of biodiversity research, the location of Riedl, 1966, and references therein). Moreover, some the Maltese Islands is strategic as they lie astride the caves are characterised by unique features because of connection between the western basin, which has gen- their physical structure and location, such as for ex- erally been very well studied, and the eastern one, for ample the `3PP Cave' in France, where true bathyal which there are only fragmented data. Located close and bathyo-abyssal organisms are able to thrive due to to the narrowest part of the channel between Africa and abnormally low and constant temperatures (Harmelin & Europe, the Maltese Archipelago occupies a key location Vacelet, 1997, and references therein). for the study of movement of flora and fauna between In recent decades, systematic surveys of dark lit- the western and eastern Mediterranean. Moreover, the toral submarine caves have received particular attention archipelago lies on the main route of the westward range from the scientific community (e.g. Bakran-Petricioli extension of Erythraean (= Lessepsian) immigrants and et al., 2007; Bianchi, Cattaneo-Vietti, Cinelli, Morri of thermophilic indigenous Mediterranean species (Le- & Pansini, 1996; Bussotti, Denitto, Guidetti & Bel- jeusne, Chevaldonn´e,Pergent-Martini, Boudouresque & monte, 2002; Boxshall & Jaume, 2000; Chevaldonn´e& P´erez,2010), and that of the eastward extension of spe- Lejeusne, 2003; Harmelin & Vacelet, 1997; Hart Jr., cies entering from the Atlantic (Sciberras & Schembri, Manning & Iliffe, 1985; Iliffe, Hart & Manning, 1985; 2007). Consequently, the Islands are ideally situated Vacelet & Boury-Esnault, 1995; Ereskovsky, Kovtun & for understanding the processes underlying the evolu- Pronin, 2015). The particular environmental conditions tion of Mediterranean biodiversity at different spatial of these habitats, including the absence of light, oligo- and temporal scales and for monitoring of the changing trophy, and reduced hydrodynamic action, make dark biogeography of the central Mediterranean. The latter is submarine caves enclave mesocosms of the deep aphotic necessary as surface hydrographical conditions change, zone in shallow coastal areas (Harmelin et al., 1985). In such as for example the ongoing northward displacement this respect, marine caves mimic the deep sea. Marine of the `15 ◦C divide' that is allowing west to east and east caves therefore conveniently serve as natural laborat- to west passage of thermophilic species (Bianchi et al., ories, and render unique opportunities for researchers 2012). to access organisms and processes that are otherwise The Maltese Islands, being almost entirely composed very difficult to study (Bakran-Petricioli et al., 2007; of limestones of Oligo-Miocene age, have an abundance Calado, Chevaldonn´e& dos Santos, 2004; Harmelin & of partially submerged (hereafter `emergent') and sub- Vacelet, 1997; Janssen, Chevaldonn´e& Arbizu, 2013; merged marine caves with different geomorphological Vacelet, Boury-Esnault & Harmelin, 1994). Further- characteristics. Caves may arise by the direct action more, marine caves harbour rare species that are unique of the sea on the limestone rock at sea-level, where the to such habitats. Due to their relatively small size and force of the waves develops fissures in the rock into clefts, ease of accessibility, environmental stability, and pres- which eventually become caves and tunnels. This ef- ence of communities of endemic and specialized species fect is enhanced if the water carries abrasive suspended (Harmelin et al., 1985), dark submarine caves are ex- material. Other caves originated on land due to karst- cellent model habitats to address important ecological fluvial processes and then became totally or partially and evolutionary questions such as the influence of life submerged due to changes in sea-level or due to tectonic cycle and habitat fragmentation on gene flow (Cheval- movement. Some such caves continue above
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