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The coast of the Gulf of Orosei (Sardinia, Italy) consists of impressive cliffs set up on dolostones and limestones characterized by wide karst systems connected to the sea. Marine caves, which are part of these system flooded by seawater through marine entrances, may be considered as extreme environments because of wide spatial and temporal environmental variability due to changing marine and terrestrial contributions. This study presents the results of the third survey carried out in summer 2016 in the Bue Marino cave, as part of a research project started in 2014 aimed at the application of Benthic Foraminifera (BF) as ecological indicators in Mediterranean marine caves for the identification of different habitats and their environmental interpretation. Sediment and water samples were collected from a total of 25 stations from two distinct sectors of the cave (North Branch and Middle Branch); sediments were analysed for living and dead BF and grain size, while Temperature, Salinity, pH and Dissolved Oxygen were measured in water samples collected close to sediment water interface. Two main foraminiferal assemblages, with distinct characteristics with respect to the typical Mediterranean shallow-water ones, were recognized by means of Hierarchical Cluster Analysis and Non-metric Multidimensional Scaling, and a Canonical Correspondence Analysis deduced their environmental significance. A well oxygenated, less saline environment with coarse bottom sediment, correlated with a mixed calcareous-agglutinated assemblage (Gavelinopsis praegeri, Rosalina spp., Eggerelloides advenus and Reophax dentaliniformis) with high species diversity (H-index 2.32–3.57) and low foraminiferal density, was exclusive of the North Branch. A scarcely oxygenated, more saline environment with fine bottom sediment enriched in vegetal debris was related to a prevalently agglutinated assemblage characterized by low species diversity (H-index 1.60–2.68), with high dominance of E. advenus (up to 83.6%) associated to Ammonia tepida, and high foraminiferal density, recognized in the Middle Branch. These different environments were interpreted considering the different modes of feeding the karst systems of the two branches. They also corresponded to two distinct ecozones, Entrance and Confluence, already recognized in earlier studies. The environmental significance of the foraminiferal ecozones recognized in this study and their comparison with the ones identified in the previous years, helped to consider the ecological zonation as a tool for detecting seasonal and, possibly, long term annual environmental variability in the marine system.

Although sponges constitute the dominant sessile organisms in marine caves, their functional role as ecosystem engineers has received little attention in this habitat type. In this study the associated macrofauna of the massive/tubular ecosystem-engineering sponges Agelas oroides and Aplysina aerophoba was studied across distinct ecological zones of two eastern Mediterranean caves. Our results revealed that the examined sponges supported a considerable associated macrofauna. A total of 86 associated taxa were found, including species reported for the first time as sponge symbionts and typical cave dwellers. Crustaceans predominated in terms of abundance but polychaetes showed the highest number of taxa. A clear differentiation was observed in the structure of the associated assemblage between the two sponges, attributed not only to the sponge species but also to differences in the surrounding environment. Density, diversity and the trophic structure of the sponge-associated macrofauna did not vary significantly along the horizontal axis of the surveyed caves. These findings suggest that sponges form a quite stable habitat, maintaining their functional role as ecosystem engineers across the studied marine caves and increasing habitat complexity in the impoverished inner dark cave sectors. A pesar de que las esponjas constituyen el grupo sésil dominante en cuevas submarinas, su papel como ingenieros del ecosistema dentro de estos ambientes ha recibido muy poca atención. En el presente trabajo, se ha estudiado la macrofauna asociada a las especies Agelas oroides y Aplysina aerophoba (ambas con una morfología masiva-tubular y consideradas como especies ingenieras del ecosistema) a lo largo de diferentes ambientes situados en dos cuevas marinas del Mediterráneo oriental. Nuestros resultados ponen de manifiesto que las dos especies de esponjas consideradas albergan una abundante fauna asociada. Se encontró un total de 86 taxones diferentes, muchos de los cuales son citados por primera vez como simbiontes de esponjas y habitantes de cuevas submarinas. Aunque los crustáceos fueron el grupo dominante en términos de abundancia, fueron los poliquetos los que presentaron la mayor riqueza taxonómica. Las comunidades asociadas a una y otra esponja presentaron claras diferencias, lo cual se atribuye no solo a diferencias relacionadas con las especies hospedadoras sino también al ambiente circundante. La densidad, diversidad y estructura trófica de la macrofauna asociada a las esponjas no varió significativamente a lo largo del eje horizontal de las cuevas muestreadas. Estos resultados sugieren que las esponjas forman un hábitat bastante estable, manteniendo su papel funcional como ingenieros del ecosistema a lo largo de las cuevas submarinas y, por tanto, incrementando la complejidad del hábitat en los sectores más internos, empobrecidos y oscuros de las cuevas.

Marine caves possess unique biocoenotic and ecological characteristics. Sessile benthic species such as sponges associated with cave habitats typically show a marked zonation from the cave entrance towards the end of the cave. We describe three semi-submerged karstic caves of 50 to 83 m length and 936 to 2,291 m3volume from the poorly explored cavernicolous fauna of North-East Bulgaria. We surveyed sponge diversity and spatial variability. Eight demosponge species were identified based on morphological and molecular data, of which six are known from the adjacent open sea waters of the Black Sea. Two species,Protosuberites denhartogivan Soest & de Kluijver, 2003 andHalichondria bowerbankiBurton, 1930, are reported from the Black Sea for the first time. The spatial sponge distribution inside the caves is in general similar, but shows some differences in species composition and distribution depending on cave relief and hydrodynamics. The species composition of sponges of Bulgarian caves is found to be different from Crimean caves. An updated checklist of the Black Sea sponges is provided.

The main objectives of this study are the establishment of a detailed description of five semi-submerged and shallow-water marine caves from the Tarkhankut Peninsula (Crimea), their biological characteristics with particular attention to species composition, and the distribution of sponge assemblages in these caves. Three semi-submerged and two submerged caves with lengths of 9–131 m and volumes of 61–3060 m3 have been investigated. All of them are karst-abrasive or karst in origin. In the investigated caves, we inventoried seven sponge species. All were recorded species of Porifera belong to the class Demospongiae and have previously been recorded also in adjacent open sea waters. These species are tolerant to different hydrological conditions, mostly temperature and salinity. Some of them have wide geographic distribution. The species composition of sponges from the shallow water caves of Crimea is quite different from the sponge composition in Mediterranean caves. This could be due to the geographic isolation of the Black Sea and the differences in the hydro-chemical parameters of the milieu (water salinity in Tarkhankut is 18–21‰).

Numerous studies have been carried out on microbialite/metazoan associations and their relative role in forming bioconstructions. It has often been suggested that microbial carbonate acts as stabilizing element of the primary skeletal framework. Here we describe unusual bioconstruction occurring in submerged marine caves. They are randomly distributed within the caves, and show different sizes and morphologies, as well as surface roughness/smoothness. These bioconstructions, named biostalactites, are constituted by a few cm-thick crust of serpulids and other metazoans associated with carbonate micrites. All the identified sessile faunas largely consist of cryptic and sciaphilic organisms indicating cave conditions. Biostalactites are the result of the interaction between metazoans and microbial activity. Their distinctive vertical or oblique downward growth from the ceiling and walls of the caves reflects the skeletal/microbial relationship. The skeletal framework produced mainly by serpulids is syndepositionally stabilized by autochthonous microbial induced micrite.

Marine caves are unique and vulnerable habitats exhibiting high biodiversity and heterogeneity, but threatened by multiple global and local disturbances. Marine caves, although widely distributed along the Mediterranean coast, suffer for the lack of quantitative data on their structure and function, which hinder their conservation status assessment. Thanks to the availability of a nearly 30-year-long series of data (1986-2013), we evaluated ecosystem change in the Bergeggi marine cave (Ligurian Sea, NW Mediterranean), a cave with a complex shape and high habitat heterogeneity. Non-taxonomic descriptors were adopted, namely growth forms (GF) and trophic guilds (TG), which are informative about ecosystem structure and functioning, respectively. The cave experienced a general trend of change during the last three decades, mainly due to the decline in the cover of sessile organisms (especially 3-dimensional forms) matched by an increase of turf and sediment, thus causing the structural and functional homogenization of the cave community. While change before 2004 had been attributed to climatic factors (especially to the summer heat waves of 1999 and 2003), the most important rate of change was observed between 2009 and 2013, coinciding with recent major beach nourishments and the extension of the neighbouring Vado Ligure harbour, thus providing evidences on the importance of local disturbances deriving from coastal interventions. Monitoring the status of cave ecosystems is urgently needed, and the use of effective indicators, such as the specific traits here adopted (morphology and feeding strategy), could provide effective tools to assist marine cave conservation.

This is the first paper documenting research on a selection of marine caves located along the coast of Capo Milazzo in the southern Tyrrhenian Sea. Three submarine and one semi-submerged caves were surveyed and sampled using underwater photo sampling. Surveys have only taken into account the sessile species belonging to the main taxa: Porifera, Anthozoa, Bryozoa and Polychaeta. Diversity indices and abundances were calculated for three sections within each explored cave: the Entrance Zone, Intermediate Zone and Bottom Zone. The richest group was Porifera with 21 taxa, followed by cnidarians, (Anthozoa), with 8 taxa, Polychaeta (5 taxa), and Bryozoa (5 taxa). Among Porifera, the presence of Petrobiona massiliana, a protected species according to SPA/BIO Protocol and the Bern Convention, must be highlighted. The encrusting forms were dominant in the Bottom Zone, the massive forms in the Intermediate Zone and the arborescent forms in the Entrance Zone. Generally, the percentage coverage of each morphological group showed a decline in the Intermediate Zone and a general increase in the Dark Zone within each cave. The S, H’ and J values showed different trends in the five caves. These differences, also evidenced by Permanova analysis, depend on the topographic specificity of each cave which, in turn, affects the gradients of the biotic and abiotic parameters. Finally, no horizontal gradient of rarefaction of the benthic sessile fauna has been detected. This study represents an important step for the management and conservation practices of these fragile ecosystems, especially in view of the forthcoming establishment of the Marine Protected Area.

The present study explores the amphipod assemblages of six marine caves on the Mediterranean coast of southern Spain. Replicate samples were taken both inside and outside each marine cave in order to characterize the amphipod fauna and the physicochemical properties of the sediment. As a result, 44 amphipods species were identified. The high number of species found in a relatively limited area highlighted the richness of the Alboran Sea fauna, which is mainly due to the mixture of species from different biogeographic areas. Harpinia genus was the dominant amphipod taxa inside marine caves and Perioculodes longimanus was also very abundant in the caverns. On the other hand, external communities were dominated by Siphonoecetes sabatieri, Metaphoxus fultoni and Photis longipes. There was a high degree of variability in both the internal and the external stations. Only the external station situated at low-medium depth showed a relatively homogeneous amphipod assemblage. The structure and behaviour of soft-bottom communities inside marine caves is difficult to predict because their environmental conditions depend on a particular combination of factors such as topography, depth and orientation. Therefore, no constant patterns were observed for species richness, Shannon diversity and abundance of amphipods in marine caves in comparison with open habitats. According to canonical correspondence analysis, sediment granulometry, organic matter and nitrogen concentration were the parameters that best explained the distribution of amphipods. Species were also classified by their tolerance to environmental pollution according to criteria followed by the Azti Marine Biotic Index and the BENTIX index. The great abundance of sensitive species at both the internal and external stations indicates the good ecological quality of the soft bottom studied. However, the suitability of biotic indices in marine caves should be tested in future studies. En el presente trabajo se estudia la fauna de anfípodos presente en el sustrato blando de seis cuevas submarinas de la costa Mediterránea del sur de España. Con el objetivo de caracterizar tanto la comunidad biótica como las características físico-químicas del sedimento, se tomaron muestras de sedimento a la salida y en el interior de las cuevas submarinas. Como resultado, se identificaron un total de cuarenta y cuatro especies de anfípodos diferentes. Este elevado número de especies encontrado en un área relativamente limitada pone de manifiesto la riqueza faunística del Mar de Alborán, la cual se atribuye principalmente a la mezcla de especies de diferentes áreas biogeográficas. El género Harpinia fue el taxón dominante en el interior de las cuevas, junto con Perioculodes longimanus. Por otro lado, la comunidad en las zonas externas estaba dominada por Siphonoecetes sabatieri, Metaphoxus fultoni y Photis longipes. Se observó un alto grado de variabilidad tanto en las estaciones internas como externas. Solo las estaciones externas situadas a profundidades bajas-medias mostraron una fauna relativamente homogénea. La estructura y el comportamiento de las comunidades que habitan los sustratos blandos en el interior de cuevas submarinas es difícil de predecir, ya que las condiciones ambientales dependen de sus particulares características en cuanto a topografía, profundidad y orientación. Por ello, no ha podido extraerse ningún patrón constante de la comparación de comunidades internas y externas para el número de especies, diversidad de Shannon o abundancia de anfípodos. La granulometría y los porcentajes de materia orgánica y nitrógeno en el sedimento fueron los parámetros que mejor explicaban la distribución de los anfípodos, de acuerdo con los resultados del análisis CCA. Las especies también fueron clasificadas en base a su tolerancia a la polución ambiental, siguiendo los criterios establecidos por los índices AMBI y BENTIX. La gran abundancia de especies sensibles refleja el buen estado ecológico de los sedimentos estudiados. Sin embargo, la aplicabilidad de los índices bióticos en cuevas submarinas debería ser testada en futuros estudios. Ministerio de Economía y Competitividad CGL2011-22474 Junta de Andalucía P11-RNM-7041

Mobile epifauna is an essential component of rocky reef ecosystems. In spite of this and the great scientific interest that the study of marine caves has aroused in the last decades, lit- tle research has been conducted on the macrofauna associated with animal substrates in subma- rine caves. This study explores the main differences between marine caves and open habitats in terms of species composition and diversity patterns of epifaunal communities in 4 different shal- low marine caves in southern Spain. Colonies of Eudendrium sp., a marine hydroid widely distrib- uted in the Mediterranean Sea, were taken from inside and outside each cave, and all associated mobile fauna were sorted and identified. More than 90% of organisms were crustaceans, with Amphipoda as the dominant group. Although the main species did not vary significantly in abun- dance between open and cave habitats, multivariate analysis carried out for the entire amphipod community showed significant differences between these 2 habitats. nMDS analysis showed that marine cave assemblages were also characterized by a higher degree of individuality, and uni- variate analysis showed a decrease in Shannon diversity and species richness with distance into the caves, a consistent pattern for all caves studied. Although the possible role of predation pres- sure in the structure of such assemblages was also discussed, we propose that the absence of plant substrates inside the caves, in conjunction with oligotrophic conditions and low siltation, are the main factors responsible for the impoverishment of the epifaunal community. The near lack of photosynthetic activity reduced the presence of herbivorous species, while the rich detritus-feeder community supported by the animal substrates was limited by the low rate of sedimentation.

The benthic macroalgal flora from the eastern Ionian coastal area of the Peninsula Salentina is scarcely studied. This study gives a contribution to the knowledge on its biodiversity in this area, which also includes marine caves, and the geographical distribution of some interesting species. A total of 174 macroalgae (119 Rhodophyta, 27 Ochrophyta, and 28 Chlorophyta) were identified. Six species are first records for the region, one of which, Liagora ceranoides, represents a new record for the Italian flora. The vegetation of most of the wave-exposed rocky substrata, as well as of the two sulphureous caves at Santa Cesarea Terme is characterised by extensive populations of Corallinales. The chorological spectrum of the flora shows a high occurrence of Indo-Pacific and Circumtropical elements, thus resulting more similar to that of floras of the Greek Ionian Sea. La flora bentónica marina de la zona costera jónica de la península Salentina, situada en el extremo sudeste de la región italiana de Apulia, está poco estudiada. Este trabajo contribuye al conocimiento de la biodiversidad de algas de esta zona e incluye también cuevas marinas, así como la distribución geográfica de algunas de sus especies más interesantes. Se han encontrado un total de 174 macroalgas (119 Rhodophyta, 27 Ochrophyta y 28 Chlorophyta). Seis especies han sido identificadas por primera vez en la región, una de las cuales, Liagora ceranoides, representa un nuevo registro para la flora italiana. La vegetación característica de la mayoría de los sustratos rocosos expuestos, así como de las dos cuevas sulfurosas de Santa Cesarea Terme, consiste en extensas poblaciones de Corallinales. El espectro corológico de la flora muestra una alta incidencia de elementos indo-pacíficos y circuntropicales, lo que la asimila a las floras del Mar Jónico griego.

Although sponges constitute the dominant sessile organisms in marine caves, their functional role as ecosystem engineers has received little attention in this habitat type. In this study the associated macrofauna of the massive/tubular ecosystem-engineering sponges Agelas oroides and Aplysina aerophoba was studied across distinct ecological zones of two eastern Mediterranean caves. Our results revealed that the examined sponges supported a considerable associated macrofauna. A total of 86 associated taxa were found, including species reported for the first time as sponge symbionts and typical cave dwellers. Crustaceans predominated in terms of abundance but polychaetes showed the highest number of taxa. A clear differentiation was observed in the structure of the associated assemblage between the two sponges, attributed not only to the sponge species but also to differences in the surrounding environment. Density, diversity and the trophic structure of the sponge-associated macrofauna did not vary significantly along the horizontal axis of the surveyed caves. These findings suggest that sponges form a quite stable habitat, maintaining their functional role as ecosystem engineers across the studied marine caves and increasing habitat complexity in the impoverished inner dark cave sectors., A pesar de que las esponjas constituyen el grupo sésil dominante en cuevas submarinas, su papel como ingenieros del ecosistema dentro de estos ambientes ha recibido muy poca atención. En el presente trabajo, se ha estudiado la macrofauna asociada a las especies Agelas oroides y Aplysina aerophoba (ambas con una morfología masiva-tubular y consideradas como especies ingenieras del ecosistema) a lo largo de diferentes ambientes situados en dos cuevas marinas del Mediterráneo oriental. Nuestros resultados ponen de manifiesto que las dos especies de esponjas consideradas albergan una abundante fauna asociada. Se encontró un total de 86 taxones diferentes, muchos de los cuales son citados por primera vez como simbiontes de esponjas y habitantes de cuevas submarinas. Aunque los crustáceos fueron el grupo dominante en términos de abundancia, fueron los poliquetos los que presentaron la mayor riqueza taxonómica. Las comunidades asociadas a una y otra esponja presentaron claras diferencias, lo cual se atribuye no solo a diferencias relacionadas con las especies hospedadoras sino también al ambiente circundante. La densidad, diversidad y estructura trófica de la macrofauna asociada a las esponjas no varió significativamente a lo largo del eje horizontal de las cuevas muestreadas. Estos resultados sugieren que las esponjas forman un hábitat bastante estable, manteniendo su papel funcional como ingenieros del ecosistema a lo largo de las cuevas submarinas y, por tanto, incrementando la complejidad del hábitat en los sectores más internos, empobrecidos y oscuros de las cuevas.

Scientific exploration of marine cave environments and anchialine ecosystems over recent decades has led to outstanding discoveries of novel taxa, increasing our knowledge of biodiversity. However, biological research on underwater caves has taken place only in a few areas of the world and relevant information remains fragmented in isolated publications and databases. This fragmentation makes assessing the conservation status of marine cave species especially problematic, and this issue should be addressed urgently given the stresses resulting from planned and rampant development in the coastal zone worldwide. The goal of the World Register of marine Cave Species (WoRCS) initiative is to create a comprehensive taxonomic and ecological database of known species from marine caves and anchialine systems worldwide and to present this as a Thematic Species Database (TSD) of the World Register of marine Species (WoRMS). WoRCS will incorporate ecological data (e.g., type of environment, salinity regimes, and cave zone) as well as geographical information on the distribution of species in cave and anchialine environments. Biodiversity data will be progressively assembled from individual database sources at regional, national or local levels, as well as from literature sources (estimate: >20,000 existing records of cave-dwelling species scattered in several databases). Information will be organized in the WoRCS database following a standard glossary based on existing terminology. Cave-related information will be managed by the WoRCS thematic editors with all data dynamically linked to WoRMS and its team of taxonomic editors. In order to mobilize data into global biogeographic databases, a Gazetteer of the Marine and Anchialine Caves of the World will be established. The presence records of species could be eventually georeferenced for submission to the Ocean Biogeographic Information System (OBIS) and constitute an important dataset for biogeographical and climate change studies on marine caves and anchialine systems.

Scientific exploration of marine cave environments and anchialine ecosystems over recent decades has led to outstanding discoveries of novel taxa, increasing our knowledge of biodiversity. However, biological research on underwater caves has taken place only in a few areas of the world and relevant information remains fragmented in isolated publications and databases. This fragmentation makes assessing the conservation status of marine cave species especially problematic, and this issue should be addressed urgently given the stresses resulting from planned and rampant development in the coastal zone worldwide. The goal of the World Register of marine Cave Species (WoRCS) initiative is to create a comprehensive taxonomic and ecological database of known species from marine caves and anchialine systems worldwide and to present this as a Thematic Species Database (TSD) of the World Register of marine Species (WoRMS). WoRCS will incorporate ecological data (e.g., type of environment, salinity regimes, and cave zone) as well as geographical information on the distribution of species in cave and anchialine environments. Biodiversity data will be progressively assembled from individual database sources at regional, national or local levels, as well as from literature sources (estimate: >20,000 existing records of cave-dwelling species scattered in several databases). Information will be organized in the WoRCS database following a standard glossary based on existing terminology. Cave-related information will be managed by the WoRCS thematic editors with all data dynamically linked to WoRMS and its team of taxonomic editors. In order to mobilize data into global biogeographic databases, a Gazetteer of the Marine and Anchialine Caves of the World will be established. The presence records of species could be eventually georeferenced for submission to the Ocean Biogeographic Information System (OBIS) and constitute an important dataset for biogeographical and climate change studies on marine caves and anchialine systems.

We investigated how large populations of several mysid species can coexist in oligo - trophic underwater marine caves and their relationships in the marine cave food web using car- bon and nitrogen stable isotopes. Isotopic signatures indicate food partitioning among the 5 spe- cies of cave-dwelling mysids from the northwest Mediterranean Sea we studied. Hemimysis speluncola feeds mainly on phytoplankton and zooplankton from outside the caves, Siriella gra- cilipes on sedimentary organic matter and zooplankton from outside, Harmelinella mariannae on small cave-dwelling crustaceans, and Hemimysis margalefi and Hemimysis lamornae mediter- ranea on sedimentary particulate organic matter. These differences in diet could promote coexis- tence of such diverse and abundant mysid faunas in marine caves by reducing interspecific com- petition for scarce resources. The analysis of both seston and cave sediments revealed that the quantity and quality of organic matter are strongly reduced in marine caves, suggesting that cave- dwelling mysids find most of their food elsewhere. This inference agrees with documented distri- butions of some of these mysids outside of caves at night where they can feed in the rich littoral zone. These migrations of some species make cave-dwelling mysids important vectors of organic matter transfer from the outside euphotic littoral zone to various locations inside caves. Outside organic matter is then made available to other cave dwellers through mysid fecal pellet production and predation, as suggested by the isotopic compositions of predators like teleost fishes, decapod crustaceans, and carnivorous cladorhizid sponges.

Submarine caves are environments of great ecological interest because of the occurrence of peculiar conditions, such as the attenuation of light and reduced water turnover, which can determine oligotrophic conditions from the entrance to the interior part of the cave. These environmental gradients may influence the distribution of the communities inhabiting submarine caves. In this study we investigated the meiofaunal community and nematode assemblages from the sediments inside and outside two submarine caves in Ustica Island Marine Protected Area (southwest Italy): Grotta Falconiera and Grotta dei Gamberi. Consistently with a general pattern of distribution reported by several studies on benthic organisms, our results showed a decrease in the abundance and changes in the taxa composition of the meiofaunal community along the exterior-interior axis of the caves, also highlighting the dissimilarity between the dark and semi-dark communities. We found a significant influence of the availability of organic matter (i.e. phytopigment concentrations) on the distribution and composition of both the meiofauna and the nematode community inside the caves. Different nematode assemblages characterized the inside and the outside of the two caves, with species occurring exclusively in the sediment of both caves, particularly in the dark portions, and completely absent in the external sediments. Environmental features of submarine caves may affect food resources inside the caves and consequently trophic nematode assemblages. Our results showed a difference in feeding strategies between nematodes inhabiting the caves and those living outside, suggesting that in the two caves investigated, bacteria might represent the most important food source for nematodes.

Scientific information on the biodiversity of marine caves in the eastern Mediterranean is limited, especially when considering the extensively studied caves of the north-western and central Mediterranean. Aiming to enhance current knowledge regarding cave communities, this study represents a first assessment of the marine cave biota of the eastern Mediterranean, as this has been defined by the European Union’s Marine Strategy Framework Directive (MSFD). Information retrieved from an extensive overview of relevant scientific documents was combined with original data recorded from 23 marine caves of the north-eastern Mediterranean. Our results report a total of 520 taxa recorded in eastern Mediterranean marine caves to date, the majority of which are sponges, polychaetes, rhodophytes, bivalves, fishes, and gastropods. These include several protected, endemic, and alien species. However, not all taxonomic groups have been equally studied among different areas and future studies are expected to raise the number of endemic and alien species. The overall observed trend is that the reported species number is generally related to sampling effort and scientific expertise. The most well-studied marine cave communities of the eastern Mediterranean are those of the Aegean Sea (especially its northern sector), which presented the highest number of species, followed by those of the Levantine. Furthermore, our research in Aegean caves revealed numerous new records for the marine cave fauna of the eastern basin, while several species are reported for the first time in the marine cave habitat. The critical need for further scientific research, monitoring, and conservation of this unique ecosystem was highlighted by (i) the presence of certain species endemic to the eastern Mediterranean coupled with a high proportion of alien species, especially in the Levantine, and (ii) the marine cave habitat availability in isolated insular areas of the eastern Mediterranean.

In spite of the high importance of mobile epifauna in all marine habitats, their pat terns of colonization and succession in marine caves have not been studied until now. In the pres ent study, we used artificial substrates deployed at 4 positions along an environmental gradient of a ~100 m long cave system, and retrieved at different times (1.5, 3 and 6 mo), to explore the changes in abundance, species richness, and community structure of the epifauna. All inner cave stations showed significantly lower species richness and abundance. Despite different dispersal modes, the dominant species detected were able to quickly (

Marine caves are vulnerable ecosystems of special conservation interest. These poorly investigated habitats support rich assemblages, including fragile long-living species as well as several protected, rare and bathyal faunal elements. Due to their rich biological diversity and complex geomorphology marine caves are important SCUBA diving hotspots. Internationally, there are various models of management of visitation and conservation of marine caves. The aim of this study was to present a preliminary comparison of different protocols for marine cave visitation, and to examine case studies from marine regions worldwide. Although marine cave habitats have been included in international environmental directives and conservation action plans (e.g. EU and Mediterranean countries) in most countries no specific regulations or management plans are implemented, even for caves located in Marine Protected Areas (MPAs). Surprisingly, few MPAs (e.g. Rowley Shoals Marine Park in Australia and National Park of Svalbard in Norway) have set specific limitations with regard to divers’ visitation levels in their caves (e.g. annual number of divers). In some cases special official permits are prerequisite for divers to visit marine caves located in protected areas (e.g. Ecological Network of the Republic of Croatia) ; though this regulation is not strictly followed for several caves. The large number of marine caves in coastal areas with extensive rocky outcrops (e.g. eastern Adriatic Sea, Aegean islands) coupled with the fact that few attempts of detailed mapping of their geographical location have been made make it practically impossible to manage human activities (e.g. SCUBA diving, spear fishing, tourist boat tours) and evaluate possible impacts on cave biota. Therefore numerous marine caves are currently exposed to unregulated visitation levels and potentially deleterious practices (e.g. souvenir collection, damage to sessile marine organisms).

The present study explores the amphipod assemblages of six marine caves on the Mediterranean coast of southern Spain. Replicate samples were taken both inside and outside each marine cave in order to characterize the amphipod fauna and the physicochemical properties of the sediment. As a result, 44 amphipods species were identified. The high number of species found in a relatively limited area highlighted the richness of the Alboran Sea fauna, which is mainly due to the mixture of species from different biogeographic areas. Harpinia genus was the dominant amphipod taxa inside marine caves and Perioculodes longimanus was also very abundant in the caverns. On the other hand, external communities were dominated by Siphonoecetes sabatieri, Metaphoxus fultoni and Photis longipes. There was a high degree of variability in both the internal and the external stations. Only the external station situated at low-medium depth showed a relatively homogeneous amphipod assemblage. The structure and behaviour of soft-bottom communities inside marine caves is difficult to predict because their environmental conditions depend on a particular combination of factors such as topography, depth and orientation. Therefore, no constant patterns were observed for species richness, Shannon diversity and abundance of amphipods in marine caves in comparison with open habitats. According to canonical correspondence analysis, sediment granulometry, organic matter and nitrogen concentration were the parameters that best explained the distribution of amphipods. Species were also classified by their tolerance to environmental pollution according to criteria followed by the Azti Marine Biotic Index and the BENTIX index. The great abundance of sensitive species at both the internal and external stations indicates the good ecological quality of the soft bottom studied. However, the suitability of biotic indices in marine caves should be tested in future studies., En el presente trabajo se estudia la fauna de anfípodos presente en el sustrato blando de seis cuevas submarinas de la costa Mediterránea del sur de España. Con el objetivo de caracterizar tanto la comunidad biótica como las características físico-químicas del sedimento, se tomaron muestras de sedimento a la salida y en el interior de las cuevas submarinas. Como resultado, se identificaron un total de cuarenta y cuatro especies de anfípodos diferentes. Este elevado número de especies encontrado en un área relativamente limitada pone de manifiesto la riqueza faunística del Mar de Alborán, la cual se atribuye principalmente a la mezcla de especies de diferentes áreas biogeográficas. El género Harpinia fue el taxón dominante en el interior de las cuevas, junto con Perioculodes longimanus. Por otro lado, la comunidad en las zonas externas estaba dominada por Siphonoecetes sabatieri, Metaphoxus fultoni y Photis longipes. Se observó un alto grado de variabilidad tanto en las estaciones internas como externas. Solo las estaciones externas situadas a profundidades bajas-medias mostraron una fauna relativamente homogénea. La estructura y el comportamiento de las comunidades que habitan los sustratos blandos en el interior de cuevas submarinas es difícil de predecir, ya que las condiciones ambientales dependen de sus particulares características en cuanto a topografía, profundidad y orientación. Por ello, no ha podido extraerse ningún patrón constante de la comparación de comunidades internas y externas para el número de especies, diversidad de Shannon o abundancia de anfípodos. La granulometría y los porcentajes de materia orgánica y nitrógeno en el sedimento fueron los parámetros que mejor explicaban la distribución de los anfípodos, de acuerdo con los resultados del análisis CCA. Las especies también fueron clasificadas en base a su tolerancia a la polución ambiental, siguiendo los criterios establecidos por los índices AMBI y BENTIX. La gran abundancia de especies sensibles refleja el buen estado ecológico de los sedimentos estudiados. Sin embargo, la aplicabilidad de los índices bióticos en cuevas submarinas debería ser testada en futuros estudios., Financial support for this work was provided by the Ministerio de Economía y Competitividad (Project CGL 2011-22474, internal reference 2011-707). It was cofinanced by FEDER funds of the European Union and by the Consejería de Economía, Innovación, Ciencia y Empleo, Junta de Andalucía (Project P11-RNM-7041).

The marine cave-dwelling mysid Hemimysis margalefi is distributed over the whole Mediterranean Sea, which contrasts with the poor dispersal capabilities of this brooding species. In addition, underwater marine caves are a highly fragmented habitat which further promotes strong genetic structuring, therefore providing highly informative data on the levels of marine population connectivity across biogeographical regions. This study investigates how habitat and geography have shaped the connectivity network of this poor disperser over the entire Mediterranean Sea through the use of several mitochondrial and nuclear markers. Five deeply divergent lineages were observed among H. margalefi populations resulting from deep phylogeographical breaks, some dating back to the Oligo-Miocene. Whether looking at the intralineage or interlineage levels, H. margalefi populations present a high genetic diversity and population structuring. This study suggests that the five distinct lineages observed in H. margalefi actually correspond to as many separate cryptic taxa. The nominal species, H. margalefi sensu stricto, corresponds to the westernmost lineage here surveyed from the Alboran Sea to southeastern Italy. Typical genetic breaks such as the Almeria-Oran Front or the Siculo-Tunisian Strait do not appear to be influential on the studied loci in H. margalefi sensu stricto. Instead, population structuring appears more complex and subtle than usually found for model species with a pelagic dispersal phase. The remaining four cryptic taxa are all found in the eastern basin, but incomplete lineage sorting is suspected and speciation might still be in process. Present-day population structure of the different H. margalefi cryptic species appears to result from past vicariance events started in the Oligo-Miocene and maintained by present-day coastal topography, water circulation and habitat fragmentation. © 2014 John Wiley & Sons Ltd.