Benthic Foraminifera as an Environmental Indicators in Mediterranean Marine Caves: A Review

Marine caves are characterized by wide environmental variability for the interaction between marine and continental processes. Their conditions may be defined as extreme for inhabiting organisms due to the enclosed morphology, lack of light, and scarcity of nutrients. Therefore, it is necessary to identify reliable ecological indicators for describing and assessing environmental conditions in these habitats even more than elsewhere. This review aims to provide the state of art related to the application of benthic foraminifera as proxies in the (paleo)ecological characterization of different habitats of marine caves. Special attention was addressed to a research project focused on Mediterranean marine caves with different characteristics, such as extent, morphology, freshwater influence, salinity, sediment type, oxygenation, and organic matter supply. This review aims to illustrate the reliability of foraminifera as an ecological and paleoecological indicator in these habitats. They respond to various environmental conditions with different assemblages corresponding to a very detailed habitat partitioning. Because marine caves may be considered natural laboratories for environmental variability, the results of these studies may be interpreted in the perspective of the global variability to understand the environmental drivers of future changes in marine systems.

Rediscovery of the rare Mediterranean marine cave stenopodid shrimp Odontozona addaia Pretus, 1990, 30 years after its original description (Crustacea: Decapoda: Stenopodidea)

Thirty years after its first finding and description, the marine cave stenopodid shrimp Odontozona addaia is here reported for the second time. The new localities, particular marine caves of southern France, are more than 300 km apart from the type locality in the Balearic Islands. First live in situ photographs are provided, and the morphological intraspecific variability is detailed by comparing the new specimens to the types. DNA sequences were also obtained for comparison with other Odontozona species. Based on both morphology and molecular analysis, closest relatives of O. addaia appear to be the western Atlantic Odontozona meloi and the eastern Mediterranean Odontozona minoica, although their detailed relationships remain unresolved.

Lithistid Demosponges of Deep-Water Origin in Marine Caves of the North-Eastern Mediterranean Sea

Desmas-bearing demosponges known as lithistids have heavily silicified skeleton and occur typically in bathyal environments of warm and tropical areas but may be found in certain shallow marine caves. Here we report, for the first time two lithistid species, i.e., Neophrissospongia endoumensis, and N. cf. nana, that were earlier known from Western Mediterranean marine caves, from four marine caves in the north-eastern Mediterranean, and their congener Neophrissospongia nolitangere from deep waters (ca. 300 m) of the Aegean Sea. All marine caves, and sections within these caves, where lithistids occur, have freshwater springs. We interpret this surprising association between lithistids and freshwater input by elevated concentration of silica in water in cave sections where such springs occur, being 8–11 times higher in comparison with shallow water outside caves, and comparable to that of deep waters, that promoted lithistids’ development. One of the studied caves harbored an abundant population of N. endoumensis which formed large masses. The age estimation of these lithistids, based on known growth rate of related deep-water sponges, suggest that they could be approximately 769–909 years old in the case of the largest specimen observed, about 100 cm large. These sponges could have colonized the caves from adjacent deep-water areas not earlier than 7,000–3,000 years ago, after the last glaciation, because earlier they were emerged. High variability of spicules, especially microscleres, and underdevelopment of megascleres may be related to silicic acid concentration.

Sponges from rhodolith beds surrounding Ustica Island marine protected area (southern Tyrrhenian Sea), with a comprehensive inventory of the island sponge fauna

The sponge fauna colonizing rhodolith beds from Ustica Island marine protected area was studied. Moreover, an inventory of the sponge species present along the island’s coasts was carried out for the first time. Analysis of rhodoliths trapped in nets used by local fishermen at two sites and data obtained from underwater video images were used to identify 25 sponge taxa, 22 of them at species level. The classes Demospongiae and Homoscleromorpha were present with 24 and 1 species, respectively. Most of the specimens were small-sized and represented by thick crusts or short erect branches. Few specimens were insinuating or excavating. Furthermore, qualitative sampling was performed on Ustica’s coralligenous formations, photophilous hard substrates and in marine caves, and the results obtained were added to the literature data. The overall checklist of sponges from Ustica encompassed 97 taxa, 90 named at species level, subdivided into 6 taxa of Calcarea, 3 of Homoscleromorpha and 88 of Demospongiae. Eleven of the species were endemic to the Mediterranean Sea, mainly concentrated on rhodoliths and in marine caves. This research adds a fundamental element to the knowledge of invertebrates associated with rhodoliths, and updates the checklist of sponges colonizing Ustica’s waters, facilitating its future monitoring.

Rock sponges (lithistid Demospongiae) of the Northeast Atlantic seamounts, with description of ten new species

Background Lithistid demosponges, also known as rock sponges, are a polyphyletic group of sponges which are widely distributed. In the Northeast Atlantic (NEA), 17 species are known and the current knowledge on their distribution is mainly restricted to the Macaronesian islands. In the Mediterranean Sea, 14 species are recorded and generally found in marine caves. Methods Lithistids were sampled in nine NEA seamounts during the scientific expeditions Seamount 1 (1987) and Seamount 2 (1993) organized by the MNHN of Paris. Collected specimens were identified through the analyses of external and internal morphological characters using light and scanning electron microscopy, and compared with material from various museum collections as well as literature records. Results A total of 68 specimens were analysed and attributed to 17 species across two orders, seven families, and seven genera, representing new records of distribution. Ten of these species are new to science, viz. Neoschrammeniella inaequalis sp. nov., N. piserai sp. nov., N. pomponiae sp. nov., Discodermia arbor sp. nov., D. kellyae sp. nov., Macandrewia schusterae sp. nov., M. minima sp. nov., Exsuperantia levii sp. nov., Leiodermatium tuba sp. nov. and Siphonidium elongatus sp. nov., and are here described and illustrated. New bathymetric records were also found for D. ramifera, D. verrucosa and M. robusta. The Meteor seamount group has a higher species richness (15 species) compared to the Lusitanian seamount group (six species). The majority of the species had their distribution restricted to one seamount, and ten are only known from a single locality, but this can be a result of sample bias. Discussion The number of species shared between the seamounts and the Macaronesian islands is very reduced. The same pattern repeats between the NEA and Mediterranean Sea. This study demonstrates that NEA seamounts are ecosystems with a higher diversity of lithistids than previously thought, increasing the number of lithistids known to occur in the NEA and Mediterranean Sea from 26 to 36 species.

APPLICATION OF RAMAN SPECTROSCOPY IN COMPARISON BETWEEN CRYPTIC MICROBIALITES OF RECENT MARINE CAVES AND TRIASSIC PATCH REEFS

ABSTRACT Microbialites are common carbonate structures in cryptic niches of marine environments throughout geological time. In this research we compare the microbialites of small bioconstructions (biostalactites) of modern submarine caves of Sicily with those developed in small crypts of Carnian patch reefs of the Dolomite Mountains (Heiligkreuz Formation, Alpe di Specie) using Raman spectroscopy, a method that allows in situ determination of the organic content of microbial components. This methodology partly solves the uncertainty of geomicrobiological studies that use bulk measurements (i.e., biomarker analyses), which make it difficult to associate mineral precipitates with a specific microbial process. In the modern marine caves, the complex biotic relationships among skeletal organisms (mainly serpulids) and microbial communities produced biostalactites in which microbially induced biomineralization is the consequence of autotrophic and chemoheterotrophic bacterial activities. Sulfate-reducing bacteria, fed by metazoan organic matter, flourish in millimetric oxygen-depleted cavities of the skeletal framework, and induce autochthonous micrite deposition and early stabilization of the biostalactites. Similar processes have been interpreted to induce the deposition of the microbialites in the Upper Triassic patch reefs of the Dolomites. These small shallow water reefs, made up mainly of scleractinian corals, sponges and red algae, hold a skeletal framework rich in millimeter- to centimeter-size cavities, ideal cryptic niches for growth of microbial communities. Specific sulfate-reducing bacteria biomarkers are first identified using bulk measurements obtained by solvent extraction. The subsequent in situ characterization of organic compounds through micro-Raman spectroscopy indicates the same biogeochemical signatures of the microbialites within the cryptic cavities of the biostalactites of modern marine caves as those inside the skeletal framework of Carnian patch reefs. These data, showing the same processes in Triassic and modern cryptic microenvironments, is evidence that the microbially mediated precipitation in confined environments is a process independent of geological time that further investigation may be able to test.

Preface

A volume on Atlantic-Mediterranean deep-sea and cave sponges dedicated to Klaus Rützler and Jean Vacelet to celebrate the publication, sixty years ago, of the first works on the biodiversity of marine caves (Laborel & Vacelet 1958; Russ & Rützler 1959).

Preliminary assessment of methanogenic microbial communities in marine caves of Zakynthos Island (Ionian Sea, Greece)

Mediterranean marine caves remain largely unexplored, while particularly limited information is available about the microbial life existing in these unique environments. The present study is a preliminary assessment of the composition of the active anaerobic microbial community colonizing the walls of newly explored systems of underwater caves and small cavities in Zakynthos Island. The interior of these caves is densely coated with egg-shaped, foam-shaped and filamentous biological structures that are characterised by a strong odor of hydrogen sulfide gas. A total of twelve structures scrapped from cave rocks were subjected to anaerobic cultivation for up to 208 days. Strong to moderate methanogenesis was observed in two different types of egg-shaped structures and one foam-like structure. Interestingly, this was observed in experiments that were performed at room temperature (i.e. 25oC) which is substantially lower than those typically considered optimum for methane production (e.g. 35oC). Analysis of the 16S rRNA genes revealed a clear dominance of archaea and bacteria closely related to known methane producers and sulfate reducers, including members of the families Methanomicrobiaceae, Desulfobulbaceae, Desulfobacteraceae, Desulfuromonaceae, Campylobacteraceae, Marinifilaceae, Clostridiaceae, Incertae Sedis – Family I & II. These results show that Mediterranean marine caves can host members of archaea and bacteria with potential biotechnological interest that deserve further investigation.