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The taxonomic composition, structure, and distribution patterns of the macrobenthos associated with Corallium rubrum were studied along the coast of Taranto (Ionian Sea), together with the main features of their red coral population. Underwater video transects were performed by professional divers at three sites in correspondence with coralligenous formations at depths from 50 to 65 m. The results revealed a patchy distribution of red coral, with colonies predominantly located in cavities on sub-vertical cliffs and large boulders. Biometric analysis indicated that young colonies predominated at all sites, while older colonies were lacking, likely because of illegal harvesting. The lower density values were recorded at S1, while S2 and S3 presented higher values. A total of 76 taxa were recorded. S1, the shallowest site, showed a prevalence of calcareous algae, while S2 and S3 showed a greater abundance of filter-feeding invertebrates (Porifera and Cnidaria) with the highest presence of Porifera at S3. The results emphasize the heterogeneity of the macrobenthos together with the high vulnerability of the red coral population, highlighting the necessity of site-specific conservation strategies to contribute to the conservation and management of benthic ecosystems in the northern Ionian Sea. [ABSTRACT FROM AUTHOR]

Abstract Background Climate change has accelerated the occurrence and severity of heatwaves in the Mediterranean Sea and poses a significant threat to the octocoral species that form the foundation of marine animal forests (MAFs). As coral health intricately relies on the symbiotic relationships established between corals and microbial communities, our goal was to gain a deeper understanding of the role of bacteria in the observed tissue loss of key octocoral species following the unprecedented heatwaves in 2022. Results Using amplicon sequencing and taxon-specific qPCR analyses, we unexpectedly found that the absolute abundance of the major bacterial symbionts, Spirochaetaceae (C. rubrum) and Endozoicomonas (P. clavata), remained, in most cases, unchanged between colonies with 0% and 90% tissue loss. These results suggest that the impairment of coral health was not due to the loss of the main bacterial symbionts. However, we observed a significant increase in the total abundance of bacterial opportunists, including putative pathogens such as Vibrio, which was not evident when only their relative abundance was considered. In addition, there was no clear relation between bacterial symbiont loss and the intensity of thermal stress, suggesting that factors other than temperature may have influenced the differential response of octocoral microbiomes at different sampling sites. Conclusions Our results indicate that tissue loss in octocorals is not directly caused by the decline of the main bacterial symbionts but by the proliferation of opportunistic and pathogenic bacteria. Our findings thus underscore the significance of considering both relative and absolute quantification approaches when evaluating the impact of stressors on coral microbiome as the relative quantification does not accurately depict the actual changes in the microbiome. Consequently, this research enhances our comprehension of the intricate interplay between host organisms, their microbiomes, and environmental stressors, while offering valuable insights into the ecological implications of heatwaves on marine animal forests. Video Abstract

The taxonomic composition, structure, and distribution patterns of the macrobenthos associated with Corallium rubrum were studied along the coast of Taranto (Ionian Sea), together with the main features of their red coral population. Underwater video transects were performed by professional divers at three sites in correspondence with coralligenous formations at depths from 50 to 65 m. The results revealed a patchy distribution of red coral, with colonies predominantly located in cavities on sub-vertical cliffs and large boulders. Biometric analysis indicated that young colonies predominated at all sites, while older colonies were lacking, likely because of illegal harvesting. The lower density values were recorded at S1, while S2 and S3 presented higher values. A total of 76 taxa were recorded. S1, the shallowest site, showed a prevalence of calcareous algae, while S2 and S3 showed a greater abundance of filter-feeding invertebrates (Porifera and Cnidaria) with the highest presence of Porifera at S3. The results emphasize the heterogeneity of the macrobenthos together with the high vulnerability of the red coral population, highlighting the necessity of site-specific conservation strategies to contribute to the conservation and management of benthic ecosystems in the northern Ionian Sea.

Abstract Marine heat waves (MHWs) have increased in frequency and intensity worldwide, causing mass mortality of benthic organisms and loss of biodiversity in shallow waters. The Mediterranean Sea is no exception, with shallow populations of habitat-forming octocorals facing the threat of local extinction. The mesophotic zone, which is less affected by MHWs, may be of ecological importance in conservation strategies for these species. However, our understanding of the response of mesophotic octocoral holobionts to changes in seawater temperature remains limited. To address this knowledge gap, we conducted a study on an iconic Mediterranean octocoral, the red coral Corallium rubrum sampled at 60 m depth and 15 °C. We exposed the colonies to temperatures they occasionally experience (18 °C) and temperatures that could occur at the end of the century if global warming continues (21 °C). We also tested their response to extremely cold and warm temperatures (12 °C and 24 °C). Our results show a high tolerance of C. rubrum to a two-month long exposure to temperatures ranging from 12 to 21 °C as no colony showed signs of tissue loss, reduced feeding ability, stress-induced gene expression, or disruption of host-bacterial symbioses. At 24 °C, however, we measured a sharp decrease in the relative abundance of Spirochaetaceae, which are the predominant bacterial symbionts under healthy conditions, along with a relative increase in Vibrionaceae. Tissue loss and overexpression of the tumor necrosis factor receptor 1 gene were also observed after two weeks of exposure. In light of ongoing global warming, our study helps predict the consequences of MHWs on mesophotic coralligenous reefs and the biodiversity that depends on them.

Marine heat waves (MHWs) have increased in frequency and intensity worldwide, causing mass mortality of benthic organisms and loss of biodiversity in shallow waters. The Mediterranean Sea is no exception, with shallow populations of habitat-forming octocorals facing the threat of local extinction. The mesophotic zone, which is less affected by MHWs, may be of ecological importance in conservation strategies for these species. However, our understanding of the response of mesophotic octocoral holobionts to changes in seawater temperature remains limited. To address this knowledge gap, we conducted a study on an iconic Mediterranean octocoral, the red coral Corallium rubrum sampled at 60 m depth and 15 °C. We exposed the colonies to temperatures they occasionally experience (18 °C) and temperatures that could occur at the end of the century if global warming continues (21 °C). We also tested their response to extremely cold and warm temperatures (12 °C and 24 °C). Our results show a high tolerance of C. rubrum to a two-month long exposure to temperatures ranging from 12 to 21 °C as no colony showed signs of tissue loss, reduced feeding ability, stress-induced gene expression, or disruption of host-bacterial symbioses. At 24 °C, however, we measured a sharp decrease in the relative abundance of Spirochaetaceae, which are the predominant bacterial symbionts under healthy conditions, along with a relative increase in Vibrionaceae. Tissue loss and overexpression of the tumor necrosis factor receptor 1 gene were also observed after two weeks of exposure. In light of ongoing global warming, our study helps predict the consequences of MHWs on mesophotic coralligenous reefs and the biodiversity that depends on them. [ABSTRACT FROM AUTHOR]

The sub-fossil red coral deposits of Sciacca (Sicily Channel) have attracted scientific attention for nearly 150 years. Their origin and formation have been long questioned and investigated, given the fact that they represent one of the most intriguing geobiological events ever to occur in the Mediterranean basin. Less attention was given to the paleo-community associated with the sub-fossil coral. Radiocarbon age determinations, in particular, were provided only for red coral, neglecting the possibility of understanding which species were simultaneously present in the coralline paleo-community and which was their role. The study of cemented coral rubble pieces revealed that Corallium rubrum covered the largest time interval (more than 3000 years) and was contemporary to many secondary epibionts over two millennia and to Madrepora oculata for about 500 years; this last finding suggested that an uncommon co-dominance between the two structuring species occurred in the nearby living communities. The lack of Fe–Mg deposits on the cemented coral rubbles coupled with the low bioerosion rate of the red coral skeletons by the demosponge Siphonodictyon coralliirubri (assessed through the analysis of the erosive paleo-scars) suggested that the deposits met with a rapid sediment cover-up. Moreover, for the first time, the analysis of a piece of cemented coral rubble of sub-fossil red coral coming from Sardinian waters confirmed that, albeit to a lesser extent, the conditions favouring the preservation of dead corals can occur also in different localities far from Sciacca. [ABSTRACT FROM AUTHOR]

The precious red coral (Corallium rubrum L.) represents one of the most fascinating marine species of the Mediterranean Sea. Several samples, including red coral together with its accompanying species, were found in the zoological collection of the Italian pioneer biologist Lazzaro Spallanzani (1729–1799), collected in the Messina Strait during his voyage in Sicily (1788). The study of these samples allowed the inclusion of numerous additional species in the traditional red coral facies as the large oyster Neopycnodonte cochlear, the giant barnacle Pachylasma giganteum, the mesophotic scleractinian Caryophyllia (Caryophyllia) cyathus. These specimens proved to be very useful in describing the diversity of the paleo-community including red coral, shedding light on its formation processes. In particular, some specimens are composed of red coral rubble consolidated and cemented with other carbonatic remains Probably, these peculiar specimens have a similar origin to those of the Sciacca Banks already known from the Sicily Channel. In fact, the two areas are prone to intense seismic activity that periodically causes mass mortalities of red coral from nearby rocky reliefs and the formation of biogenic detritus, while the resulting chemical environment of the water and the sediments allows the consolidation of the carbonatic remains. [ABSTRACT FROM AUTHOR]

Corals are ecosystem engineers whose tree-like structures give three-dimensional complexity to the habitat. Their population dynamics are affected by recruitment and juvenile survival. Therefore, several defense strategies, such as the formation of hard skeletons and/or spicules, have evolved to protect these vulnerable stages. The family Coralliidae, to which “precious corals” belong, represent an exception in the order Scleralcyonacea, as they form hard CaCO3 skeletons and small CaCO3 structures, the sclerites. The skeletogenesis of Corallium species is relatively well documented in adult colonies but remains poorly known in the early stages of the development of new colonies. To shed light on the timing of Corallium rubrum’s early skeleton formation and the role of sclerites, we focused on the first 4-years of life, applying different techniques, from scanning electron microscopy to synchrotron tomography and laser ablation inductively coupled plasma-mass spectrometry. Our results show that: 1) the first visible sclerites in the primary polyp appear at least 12 days after larval settlement, which is associated with a high CaCO3 production rate (4.5 ± 2.3 μg of CaCO3 per day). Furthermore, growth rings are visible in the sclerites, showing that fully matured sclerites grow fast, probably in 3 to 4 days. 2) Sclerites are the only biomineral product in the first year of life of C. rubrum’s colonies. 3) The evidence of a consolidated axial skeleton, intended as the inner part of the skeleton characteristic of the adult red coral (the medullary zone, MZ), is recorded for the first time in 2-year-old colonies. 4) The annular zone (AZ) around the medullary zone starts forming not before four years after settlement. Thus, primary polyp builds a deformable armor made of only sclerites during the first year. This shelter provides mechanical protection from abrasion and predation to early settled colonies. After two years, settlers are firmly and mineralogically attached to the substratum, which makes them less vulnerable to predation than younger recruits that are not anchored by the skeleton.

The precious coral Corallium rubrum (Linnaeus, 1758) is a charismatic Mediterranean species. A recent large-scale investigation along the Italian coast highlighted its widespread occurrence at mesophotic and upper bathyal depths, especially on coralligenous and bathyal vertical hardgrounds. The lack of morphometric data limited the considerations on the structure and health status of the populations, fundamental to identify the most vulnerable sites and the correct management actions. For this reason, a ROV dataset, consisting of 624 dives carried out between 40 m and 1825 m in the Ligurian Sea, Tyrrhenian Sea, and Sicily Channel, was analysed to extrapolate quantitative data to describe the populations. Ten random frames were obtained from each of the 170 sites hosting red coral. Density, height, and entanglement were evaluated for about 15700 colonies counted in the frames. The densest populations were mainly found between 40 m and 80 m, with a clear latitudinal density decrease. The mesophotic populations were characterized by both scattered and densely aggregated colonies, while the bathyal ones were dominated by sparse colonies. This study identified 17 major coral areas based on the geographic proximity of the sites hosting red coral and their topographic and oceanographic affinity. The size-frequency distribution of the heights was skewed towards the smaller classes in almost all populations, with a modal class between 2 cm and 4 cm. This study depicted a stress situation of the populations throughout the entire study area, which could be correlated also to the long-term harvesting pressure carried out in the basins. Two additional sources of direct mortality were pointed out in this study. Entanglement to artisanal and recreational fishing gear interested about 18% of the recorded colonies, mainly at mesophotic depths. Almost all populations suffered from mechanical entanglement, with the highest percentages in the Ligurian Sea and Sicilian areas. This study also highlighted a massive occurrence of recent deep mortality events, mainly along the eastern and southern coast of Sardinia and in the Campanian Archipelago. Thirty sites with extensive patches of dead colonies still in place were reported from 70 m to around 200 m, but their formation remains unclear.

Abstract Background The process of calcium carbonate biomineralization has arisen multiple times during metazoan evolution. In the phylum Cnidaria, biomineralization has mostly been studied in the subclass Hexacorallia (i.e. stony corals) in comparison to the subclass Octocorallia (i.e. red corals); the two diverged approximately 600 million years ago. The precious Mediterranean red coral, Corallium rubrum, is an octocorallian species, which produces two distinct high-magnesium calcite biominerals, the axial skeleton and the sclerites. In order to gain insight into the red coral biomineralization process and cnidarian biomineralization evolution, we studied the protein repertoire forming the organic matrix (OM) of its two biominerals. Results We combined High-Resolution Mass Spectrometry and transcriptome analysis to study the OM composition of the axial skeleton and the sclerites. We identified a total of 102 OM proteins, 52 are found in the two red coral biominerals with scleritin being the most abundant protein in each fraction. Contrary to reef building corals, the red coral organic matrix possesses a large number of collagen-like proteins. Agrin-like glycoproteins and proteins with sugar-binding domains are also predominant. Twenty-seven and 23 proteins were uniquely assigned to the axial skeleton and the sclerites, respectively. The inferred regulatory function of these OM proteins suggests that the difference between the two biominerals is due to the modeling of the matrix network, rather than the presence of specific structural components. At least one OM component could have been horizontally transferred from prokaryotes early during Octocorallia evolution. Conclusion Our results suggest that calcification of the red coral axial skeleton likely represents a secondary calcification of an ancestral gorgonian horny axis. In addition, the comparison with stony coral skeletomes highlighted the low proportion of similar proteins between the biomineral OMs of hexacorallian and octocorallian corals, suggesting an independent acquisition of calcification in anthozoans.

The precious red coral (Corallium rubrum L.) represents one of the most fascinating marine species of the Mediterranean Sea. Several samples, including red coral together with its accompanying species, were found in the zoological collection of the Italian pioneer biologist Lazzaro Spallanzani (1729–1799), collected in the Messina Strait during his voyage in Sicily (1788). The study of these samples allowed the inclusion of numerous additional species in the traditional red coral facies as the large oyster Neopycnodonte cochlear, the giant barnacle Pachylasma giganteum, the mesophotic scleractinian Caryophyllia (Caryophyllia) cyathus. These specimens proved to be very useful in describing the diversity of the paleo-community including red coral, shedding light on its formation processes. In particular, some specimens are composed of red coral rubble consolidated and cemented with other carbonatic remains Probably, these peculiar specimens have a similar origin to those of the Sciacca Banks already known from the Sicily Channel. In fact, the two areas are prone to intense seismic activity that periodically causes mass mortalities of red coral from nearby rocky reliefs and the formation of biogenic detritus, while the resulting chemical environment of the water and the sediments allows the consolidation of the carbonatic remains.

Imaging the marine environment is more and more useful to understand relationships between species, as well as natural processes. Developing photogrammetry allowed the use of 3D measuring to study populations dynamics of sessile organisms at various scales: from colony to population. This study focuses on red coral (Corallium rubrum), as known as precious coral. Metrics measured at a colony scale (e.g., maximum height, diameter and number of branches) allowed population understanding and a comparison between an old (Cerbère-Banyuls reserve) vs. a new (Calanques National Park) MPA. Our results suggested a 5-year time step allows the appearance of a significant difference between populations inside vs. outside the Calanques National Park no-take zones. Red coral colonies were taller and had more branches inside no-take zones. A significant difference was still observable for the populations inside the Cerbère-Banyuls reserve after 40 years of protection, reflecting the sustainability and effectiveness of precautionary measures set by the reserve. The impacts at the local level (mechanical destruction) and those presumed to occur via global change (climatic variations) underline the need to develop strategies both to follow the evolutions of red coral populations but also to understand their resilience. Photogrammetry induced modeling is a time and cost effective as well as non-invasive method which could be used to understand population dynamics at a seascape scale on coralligenous reefs.

Marine protected areas (MPAs) are one of the most efficient conservation tools to buffer marine biodiversity loss induced by human activities. Beside effective enforcement, an accurate understanding of the eco-evolutionary processes underlying the patterns of biodiversity is needed to reap the benefits of management policies. In this context, integrating population genetics with demographic data, the demo-genetic approach, is particularly relevant to shift from a “species-based pattern” toward an “eco-evolutionary-based processes” conservation. Here, targeting a key species in the Mediterranean coralligenous, the red coral, Corallium rubrum, in an emblematic Mediterranean MPA, the “Réserve Naturelle de Scandola” (France), we applied demo-genetic approaches at two contrasted spatial scales, among populations and within one population, to (i) infer the demographic connectivity among populations in the metapopulation network and (ii) shed new light on the genetic connectivity and on the demographic transitions underlying the dynamics of a near-pristine population. Integrating different spatial and temporal scales, we demonstrated (i) an apparent temporal stability in the pattern of genetic diversity and structure in the MPA in spite of a dramatic demographic decline and (ii) contrasted levels of genetic isolation but substantial demographic connectivity among populations. Focusing on the near-pristine population, we complemented the characterization of red coral demographic connectivity suggesting (iii) temporal variability and (iv) the occurrence of collective dispersal. In addition, we demonstrated (v) contrasted patterns of spatial genetic structure (SGS), depending on the considered stage-class (adults vs. juveniles), in the near-pristine population. This last result points out that the overall SGS resulted from a restricted dispersal of locally produced juveniles (SGS among adults and juveniles) combined to mortality during early life stages (decrease of SGS from juveniles to adults). Demonstrating the occurrence of two management units and the importance of two populations (CAVB and ALE) for the network of connectivity, we made recommendations for the management of the Réserve Naturelle de Scandola. Besides, we contributed to the implementation of scientifically driven restoration protocols in red coral by providing estimates for the size, density, and distances among patches of transplanted colonies.

Background: The process of calcium carbonate biomineralization has arisen multiple times during metazoan evolution. In the phylum Cnidaria, biomineralization has mostly been studied in the subclass Hexacorallia (i.e. stony corals) in comparison to the subclass Octocorallia (i.e. red corals); the two diverged approximately 600 million years ago. The precious Mediterranean red coral, Corallium rubrum, is an octocorallian species, which produces two distinct highmagnesium calcite biominerals, the axial skeleton and the sclerites. In order to gain insight into the red coral biomineralization process and cnidarian biomineralization evolution, we studied the protein repertoire forming the organic matrix (OM) of its two biominerals. Results: We combined High-Resolution Mass Spectrometry and transcriptome analysis to study the OM composition of the axial skeleton and the sclerites. We identified a total of 102 OM proteins, 52 are found in the two red coral biominerals with scleritin being the most abundant protein in each fraction. Contrary to reef building corals, the red coral organic matrix possesses a large number of collagen-like proteins. Agrin-like glycoproteins and proteins with sugar-binding domains are also predominant. Twenty-seven and 23 proteins were uniquely assigned to the axial skeleton and the sclerites, respectively. The inferred regulatory function of these OM proteins suggests that the difference between the two biominerals is due to the modeling of the matrix network, rather than the presence of specific structural components. At least one OM component could have been horizontally transferred from prokaryotes early during Octocorallia evolution. Conclusion: Our results suggest that calcification of the red coral axial skeleton likely represents a secondary calcification of an ancestral gorgonian horny axis. In addition, the comparison with stony coral skeletomes highlighted the low proportion of similar proteins between the biomineral OMs of hexacorallian and octocorallian corals, suggesting an independent acquisition of calcification in anthozoans. [ABSTRACT FROM AUTHOR]

Mediterranean red coral Corallium rubrum is considered the most precious coral worldwide. Harvesting activities are performed by licensed scuba divers and managed through the recent pan-Mediterranean management plan issued by General Fisheries Commission for the Mediterranean (GFCM) along with measures locally enacted, imposing limits on licenses, harvesting season, minimum depth of dive, and size. The use of Remotely Operated Vehicles (ROVs) is prohibited, with the only exception being for scientific purposes. Despite measures already in force, the implementation of additional management tools has been recently recommended. This article reports results from the first monitoring campaign on C. rubrum harvesting based on ROVs for seabed exploration and Onboard Scientific Observers (OSOs), carried out from 2012 to 2015 along the coast of Sardinia (Mediterranean Sea—Western basin). More than 450 dives were monitored, confirming how ROV’s support eases the scouting of exploitable banks, leading to increases in catches. OSOs reported the collection of colonies below the minimum reference size and catches/dive above limits. Onboard observers collected data also on colony diameter, which is crucial for the estimation of population size structure and exploitation status. OSOs proved to be valid tools in providing additional and reliable information on red coral harvesting, thus deserving to be included among mandatory measures for the sustainable exploitation of red coral in the Mediterranean Sea.

Corallium rubrum and Eunicella cavolini are two octocorals, reported as co-occurring species in the deep rocky habitats of the Mediterranean Sea with a high hydrodynamic and moderate eutrophication. Their spatial distribution and demography in the deep sea are mainly affected by temperature and direct and indirect anthropogenic activities; however, knowledge of the factors that potentially influence their co-existence is scarce. This paper provides novel data on the distribution and demography of these two species, at depths between 50 and 290 m in the Western Mediterranean Sea, providing insights on their co-occurrence. Both species exhibited the highest population density at deeper sites (>150 m), showing an inverse size–density relation. Density values ranged from 0.03 colonies m−2 to 32 and 80 col. m−2 for yellow gorgonian and red coral, respectively. The two species co-occurred in 13% of the total frames examined, mostly dwelling between 120 and 160 m depth. Distance-based linear modeling (DistLM) emphasized that when co-occurring the variability of the two species’ densities were significantly driven by the density—rather than the morphology (i.e., height)—of the other species. We stress the need for further studies to elucidate the possible mutual effects of suspension feeders and to test the role of different environmental factors potentially influencing inter-specific relationships.

Polyp activity in passive suspension feeders has been considered to be affected by several environmental factors such as hydrodynamics, water temperature and food concentration. To better elucidate the driving forces controlling polyp expansion in these organisms and the potential role of particle concentration, the octocoral Corallium rubrum was investigated in accordance with two approaches: (1) high-frequency in-situ observations examining various environmental and biological variables affecting the water column, and (2) video-recorded flume-controlled laboratory experiments performed under a range of environmental and biological conditions, in terms of water temperature, flow speed, chemical signals and zooplankton. In the field, C. rubrum polyp expansion correlated positively with particle (seston and zooplankton) concentration and current speed. This observation was confirmed by the flume video records of the laboratory experiments, which showed differences in polyp activity due to changes in temperature and current speed, but especially in response to increasing nutritional stimuli. The maximum activity was observed at the highest level of nutritional stimulus consisting of zooplankton. Zooplankton and water movement appeared to be the main factors controlling polyp expansion. These results suggest that the energy budget of passive suspension feeders (and probably the benthic community as a whole) may rely on their ability to maximise prey capture during food pulses. The latter, which may be described as discontinuous organic matter (dead or alive) input, may be the key to a better understanding of benthic-pelagic coupling processes and trophic impacts on animal forests composed of sessile suspension feeders.

The high-Mg calcite skeleton of Corallium rubrum was analyzed from living colonies collected from a wide range of depths (15 m to 607 m) and environmental settings in the Mediterranean Sea. An overarching goal was to better understand the calcification process and incorporation of elements into the slow-growing skeleton of both shallow and deep-water specimens, and more specifically, to clarify the veracity of geochemical proxies for reconstructing seawater temperatures. The coral internal structure including growth bands were determined by scanning electronic microscopy (SEM) and petrographic techniques. Trace elements (Li, B, Mg, Sr and Ba) compositions of the coral skeleton were obtained by solution and laser ablation inductively coupled plasma mass spectrometry (ICP-MS). Results show that the skeleton of the deep-water specimens has an internal microstructure similar to shallow-water colonies, with the medullar and the annular zones clearly distinguishable in both shallow and deep-water specimens. In general, the bands of the deep-water samples are very thin and equidistant compared to the irregular banding of the shallow-water specimens. Banding differences relate to the contrasting environmental dynamics, with the shallow-water specimens being exposed to large seasonal fluctuations compared to the relative stable conditions of those inhabiting bathyal depths. The inner medullar and outer annular portions differ in their trace element concentrations: Mg/Ca, Sr/Ca and Li/Ca ratios are higher in the medullar zone and seem to be influenced by growth kinetics, whereas B/Ca and Ba/Ca are similar in both zones and hence unaffected by growth rate. The variability of elemental ratios is lower in the deep-water specimens. Growth bands are highly correlated to Mg/Ca, Sr/Ca and Li/Ca, suggesting common mechanism(s) controlling the incorporation of these elements into the coral skeleton. Mg/Ca ratios were especially depleted in the dark bands. Although the mean Mg/Ca of each specimen is positively correlated to ambient seawater temperature, the intra-annual variations and amplitudes differ and do not correlate to the band widths. These findings suggest that intra- and inter-annual variations of Mg/Ca cannot be used to reconstruct a continuous time-series of long-term seasonal temperature records. However, the mean Mg/Ca composition can serve as a valuable proxy to estimate mean palaeoseawater temperature at a given site within the Mediterranean. • Shallow and deep-water Corallium rubrum have equivalent skeletal architecture, suggesting depth-independent growth patterns. • Mean Mg/Ca reconstructs mean temperature, while inter-annual timeseries reconstruction are hampered by vital effects. • Mg/Ca, Sr/Ca and Li/Ca from the annular zone are correlated, suggesting a common mechanism(s) of incorporation. • Mg/Ca, Sr/Ca and Li/Ca from the annular zone are correlated to radial width, while Ba and B are independent of growth rate. • Ba/Ca possibly reflects [Ba] sw with low concentrations in shallow-water and high concentrations at bathyal depth. [ABSTRACT FROM AUTHOR]

Polyp activity in passive suspension feeders has been considered to be affected by several environmental factors such as hydrodynamics, water temperature and food concentration. To better elucidate the driving forces controlling polyp expansion in these organisms and the potential role of particle concentration, the octocoral Corallium rubrum was investigated in accordance with two approaches: (1) high-frequency in-situ observations examining various environmental and biological variables affecting the water column, and (2) video-recorded flume-controlled laboratory experiments performed under a range of environmental and biological conditions, in terms of water temperature, flow speed, chemical signals and zooplankton. In the field, C. rubrum polyp expansion correlated positively with particle (seston and zooplankton) concentration and current speed. This observation was confirmed by the flume video records of the laboratory experiments, which showed differences in polyp activity due to changes in temperature and current speed, but especially in response to increasing nutritional stimuli. The maximum activity was observed at the highest level of nutritional stimulus consisting of zooplankton. Zooplankton and water movement appeared to be the main factors controlling polyp expansion. These results suggest that the energy budget of passive suspension feeders (and probably the benthic community as a whole) may rely on their ability to maximise prey capture during food pulses. The latter, which may be described as discontinuous organic matter (dead or alive) input, may be the key to a better understanding of benthic-pelagic coupling processes and trophic impacts on animal forests composed of sessile suspension feeders. [ABSTRACT FROM AUTHOR]

Global warming and overexploitation both threaten the integrity and resilience of marine ecosystems. Many calls have been made to at least partially offset climate change impacts through local conservation management. However, a mechanistic understanding of the interactions of multiple stressors is generally lacking for habitat‐forming species; preventing the development of sound conservation strategies.We examined the effectiveness of no‐take marine protected areas (MPAs) at enhancing structural complexity and resilience to climate change on populations of an overexploited and long‐lived octocoral. We used long‐term data over eight populations, subjected to varying levels of disturbances, and Integral Projection Models to understand how the overfishing and mass‐mortality events shape the stochastic dynamics of the Mediterranean red coral Corallium rubrum.Marine protected areas largely reduced colony partial mortality (i.e. shrinkage), enhancing the structural complexity of coral populations. However, there were no significant differences in individual mortality or population growth rates between protected and exploited populations. In contrast, warming had detrimental consequences for the long‐term viability of red coral populations, driving steady declines and potential local extinctions due to sharp effects in survival rates. Stochastic demographic models revealed only a weak compensatory effect of MPAs on the impacts of warming.Policy implications. Our results suggest that marine protected areas (MPAs) are an effective local conservation tool for enhancing the structural complexity of red coral populations. However, MPAs may not be enough to ensure red coral's persistence under future increases in thermal stress. Accordingly, conservation strategies aiming to ensure the persistence and functional role of red coral populations should include management actions at both local (well‐enforced MPAs) and global scales (reductions in greenhouse gas emissions). Finally, this study unravels the divergent demographic consequences that can arise from multiple stressors and highlights the key role of demography in better understanding and predicting the consequences of combined impacts for vulnerable ecosystems. [ABSTRACT FROM AUTHOR]

Background. Larval settlement and intra-specific interactions during the recruitment phase are crucial in determining the distribution and density of sessile marine populations. Marine caves are confined and stable habitats. As such, they provide a natural laboratory to study the settlement and recruitment processes in sessile invertebrates, including the valuable Mediterranean red coral Corallium rubrum. In the present study, the spatial and temporal variability of red coral settlers in an underwater cave was investigated by demographic and genetic approaches. Methods. Sixteen PVC tiles were positioned on the walls and ceiling of the Colombara Cave, Ligurian Sea, and recovered after twenty months. A total of 372 individuals of red coral belonging to two different reproductive events were recorded. Basal diameter, height, and number of polyps were measured, and seven microsatellites loci were used to evaluate the genetic relationships among individuals and the genetic structure. Results. Significant differences in the colonization rate were observed both between the two temporal cohorts and between ceiling and walls. No genetic structuring was observed between cohorts. Overall, high levels of relatedness among individuals were found. Conclusion. The results show that C. rubrumindividuals on tiles are highly related at very small spatial scales, suggesting that nearby recruits are likely to be sibs. Self-recruitment and the synchronous settlement of clouds of larvae could be possible explanations for the observed pattern. [ABSTRACT FROM AUTHOR]

Background Larval settlement and intra-specific interactions during the recruitment phase are crucial in determining the distribution and density of sessile marine populations. Marine caves are confined and stable habitats. As such, they provide a natural laboratory to study the settlement and recruitment processes in sessile invertebrates, including the valuable Mediterranean red coral Corallium rubrum. In the present study, the spatial and temporal variability of red coral settlers in an underwater cave was investigated by demographic and genetic approaches. Methods Sixteen PVC tiles were positioned on the walls and ceiling of the Colombara Cave, Ligurian Sea, and recovered after twenty months. A total of 372 individuals of red coral belonging to two different reproductive events were recorded. Basal diameter, height, and number of polyps were measured, and seven microsatellites loci were used to evaluate the genetic relationships among individuals and the genetic structure. Results Significant differences in the colonization rate were observed both between the two temporal cohorts and between ceiling and walls. No genetic structuring was observed between cohorts. Overall, high levels of relatedness among individuals were found. Conclusion The results show that C. rubrumindividuals on tiles are highly related at very small spatial scales, suggesting that nearby recruits are likely to be sibs. Self-recruitment and the synchronous settlement of clouds of larvae could be possible explanations for the observed pattern.

Background. Larval settlement and intra-specific interactions during the recruitment phase are crucial in determining the distribution and density of sessile marine populations. Marine caves are confined and stable habitats. As such, they provide a natural laboratory to study the settlement and recruitment processes in sessile invertebrates, including the valuable Mediterranean red coral Corallium rubrum. In the present study, the spatial and temporal variability of red coral settlers in an underwater cave was investigated by demographic and genetic approaches. Methods. Sixteen PVC tiles were positioned on the walls and ceiling of the Colombara Cave, Ligurian Sea, and recovered after twenty months. A total of 372 individuals of red coral belonging to two different reproductive events were recorded. Basal diameter, height, and number of polyps were measured, and seven microsatellites loci were used to evaluate the genetic relationships among individuals and the genetic structure. Results. Significant differences in the colonization rate were observed both between the two temporal cohorts and between ceiling and walls. No genetic structuring was observed between cohorts. Overall, high levels of relatedness among individuals were found. Conclusion. The results show that C. rubrum individuals on tiles are highly related at very small spatial scales, suggesting that nearby recruits are likely to be sibs. Selfrecruitment and the synchronous settlement of clouds of larvae could be possible explanations for the observed pattern. [ABSTRACT FROM AUTHOR]

Understanding the role of the environment in shaping the evolution of life histories remains a major challenge in ecology and evolution. We synthesize longevity patterns of marine sessile species and find strong positive relationships between depth and maximum lifespan across multiple sessile marine taxa, including corals, bivalves, sponges and macroalgae. Using long-term demographic data on marine sessile and terrestrial plant species, we show that extreme longevity leads to strongly dampened population dynamics. We also used detailed analyses of Mediterranean red coral, with a maximum lifespan of 532 years, to explore the life-history patterns of long-lived taxa and the vulnerability to external mortality sources that these characteristics can create. Depth-related environmental gradients--including light, food availability, temperature and disturbance intensity--drive highly predictable distributions of life histories that, in turn, have predictable ecological consequences for the dynamics of natural populations. [ABSTRACT FROM AUTHOR]

The cDNA sequence of arginine kinase (AK) from the precious coral Corallium rubrum was assembled from transcriptome sequence data, and the deduced amino acid sequence of 364 residues was shown to conserve the structural features characteristic of AK. Based on the amino acid sequence, the DNA coding C. rubrum AK was synthesized by overlap extension PCR to prepare the recombinant enzyme. The following kinetic parameters were determined for the C. rubrum enzyme: K (0.10 mM), K (0.79 mM), K (0.23 mM), K (2.16 mM), and k (74.3 s). These are comparable with the kinetic parameters of other AKs. However, phylogenetic analysis suggested that the C. rubrum AK sequence has a distinct origin from that of other known cnidarian AKs with unusual two-domain structure. Using oligomers designed from the sequence of C. rubrum AK, the coding region of genomic DNA of another coral Paracorallium japonicum AK was successfully amplified. Although the nucleotide sequences differed between the two AKs at 14 positions in the coding region, all involved synonymous substitutions, giving the identical amino acid sequence. The P. japonicum AK gene contained one intron at a unique position compared with other cnidarian AK genes. Together with the observations from phylogenetic analysis, the comparison of exon/intron organization supports the idea that two distinct AK gene lineages are present in cnidarians. The difference in the nucleotide sequence between the coding regions of C. rubrum and P. japonicum AKs was 1.28%, which is twice that (0.54%) of mitochondrial DNA, is consistent with the general observation that the mitochondrial genome evolves slower than the nuclear one in cnidarians. [ABSTRACT FROM AUTHOR]

Sexual reproduction is widespread among eukaryotes, and the sex-determining processes vary greatly among species. While genetic sex determination (GSD) has been intensively described in bilaterian species, no example has yet been recorded among non-bilaterians. However, the quasi-ubiquitous repartition of GSD among multicellular species suggests that similar evolutionary forces can promote this system, and that these forces could occur also in non-bilaterians. Studying sex determination across the range of Metazoan diversity is indeed important to understand better the evolution of this mechanism and its lability. We tested the existence of sex-linked genes in the gonochoric red coral (Corallium rubrum, Cnidaria) using restriction site-associated DNA sequencing. We analysed 27 461 single nucleotide polymorphisms (SNPs) in 354 individuals from 12 populations including 53 that were morphologically sexed. We found a strong association between the allele frequencies of 472 SNPs and the sex of individuals, suggesting an XX/XY sex-determination system. This result was confirmed by the identification of 435 male-specific loci. An independent test confirmed that the amplification of these loci enabled us to identify males with absolute certainty. This is the first demonstration of a GSD system among non-bilaterian species and a new example of its convergence in multicellular eukaryotes.

Sulfur has been recognized in biogenic calcites for a long time. However, its structural position is matter of debate. For some authors, sulfur is a marker of the organic matrix while it is part of the calcite structure itself for others. To better understand the place of sulfur in calcite, sulfated magnesian calcites (S-MgCalcite) have been synthetized at high pressure and temperature and studied by μ-XANES spectroscopy. S-MgCalcite XANES spectra show two different types of sulfur: sulfate (SO 4 2− ) as a predominant species and a small contribution of sulfite (SO 3 2− ), both substituting for carbonate ions in the calcite structure. To address the question of the position of sulfur in biogenic calcites, the oxidation states of sulfur in the skeleton and organic tissues of Corallium rubrum have been investigated by micro X-ray fluorescence (μ-XRF) and sulfur K-edge micro X-ray absorption near edge structure (μ-XANES) spectroscopy at the European Synchrotron Radiation Facility (ESRF, Grenoble, France) on beamline ID21. In the skeleton, sulfur is mainly present as oxidized sulfur SO 4 2− (+VI), plus a weak sulfite contribution. XANES spectra indicate that sulfur is inorganically incorporated as sulfur structurally substituted to carbonate ions (SSS). Although an organic matrix is present in the red coral skeleton, reduced organic sulfur could not be detected by μ-XANES spectroscopy in the skeleton probably due to low organic/inorganic sulfur ratio. In the organic tissues surrounding the skeleton, several sulfur oxidation states have been detected including disulfide (S–S), thioether (R–S–CH 3 ), sulfoxide (SO 2 ), sulfonate (SO 2 O − ) and sulfate (SO 4 2− ). The unexpected occurrence of inorganic sulfate within the organic tissues suggests the presence of pre-organized organic/inorganic complexes in the circulatory system of the red coral, precursors to biomineralization ahead of the growth front. [ABSTRACT FROM AUTHOR]

The aim of this paper is to supply an overview of all historical and recent knowledge on landings, fishing and geographic distribution of the red coral banks along the Italian coasts in order to make a contribution to the conservation and future management of this resource. Along the Italian coasts, the banks have been exploited for millennia, using non-selective trawling gear that was banned in Europe in 1994. Today, harvesting is allowed only by scuba divers and regulated by specific laws. We examined 153 years of history of coral fishing in Italy, from Unification (1861) to 2014. Data about the coralline fleets and the amount of coral landings were recorded for the considered span of time. From a quantitative point of view, the coral fishing in Italy in the last 150 years has been dominated by the sub-fossil coral reservoirs from the Sciacca Banks (Sicily Channel), where an extraordinary amount of 18,000 tons was collected in 34 years (1875–1888 and 1893–1914). This amount represents about 90% of all red coral harvested along the Italian coast in the last 150 years. Excluding this period, the average annual yield was initially around 100 tons, decreasing to 28 tons 100 years later, therefore demonstrating a severe overexploitation of the resource. The great part of the deep red coral banks was abandoned because harvesting was no longer profitable. Nevertheless, quantitative data suggest that red coral banks, even though overexploited, are still widespread along Italian coasts, mainly in shallow waters. These banks show a remarkable persistence although with a generalised shift towards smaller colony sizes and higher density. The time span necessary in recovering the pristine structure, following the end of the fishing activities, appears very long and the populations of several areas might be unable to re-colonise the old and overexploited banks. [ABSTRACT FROM AUTHOR]

Octocorals are marine modular organisms with high ecological and economic importance. Mediterranean Red Coral ( Corallium rubrum ), is endemic to the Mediterranean sea and neighboring Atlantic rocky shores and has been exploited for jewelry since ancient times. Despite the lack of photosynthetic symbionts ( Symbiodinium spp.), red coral growth and survival do depend on sea water temperature, as well as on trophic conditions and other physico-chemical parameters. We developed and applied a mechanistic numerical model to describe the growth of a C. rubrum colony (polyps number, polyp and gametes biomass, skeletal inorganic and organic matter) as a function of food availability and seawater temperature. The model follows a bioenergetic approach and is calibrated vs available experimental observations. Model results highlight that larger colonies are more sensitive to high temperature and actual limits of the ecological niche also depend on food availability, hydrodynamic condition and coral morphology. Bioenergetic considerations also support the conclusion that, though a modular organism, red coral exhibits constrained growth, because of the competition for available food between polyps from the same colony. [ABSTRACT FROM AUTHOR]

This study investigates the energetic investment during spawning of two Mediterranean gorgonians characterized by different reproductive strategies: Corallium rubrum (internal brooder) and Paramuricea clavata (surface brooder). Sexual products (number of oocytes and spermatic sacs) were quantified, and biochemical characteristics (lipid content and free fatty acid content and composition) were determined to investigate the parental energetic investment and demand in reproduction. Results suggested that the majority of the energetic cost was due to reproductive activity (i.e., gametogenesis and spawning). The two species exhibited different life history strategies, with P. clavata investing more energy in reproduction than C. rubrum. However, P. clavata is reproductively more sensitive to inter-annual changes in environmental conditions. [ABSTRACT FROM AUTHOR]

Commercially harvested since ancient times, the highly valuable red coral Corallium rubrum (Linnaeus, 1758) is an octocoral endemic to the Mediterranean Sea and adjacent Eastern Atlantic Ocean, where it occurs on rocky bottoms over a wide bathymetric range. Current knowledge is restricted to its shallow populations (15-50 m depth), with comparably little attention given to the deeper populations (50-200 m) that are nowadays the main target of exploitation. In this study, red coral distribution and population structure were assessed in three historically exploited areas (Amalfi, Ischia Island and Elba Island) in the Tyrrhenian Sea (Western Mediterranean Sea) between 50 and 130 m depth by means of ROV during a cruise carried out in the summer of 2010. Red coral populations showed a maximum patch frequency of 0.20 ± 0.04 SD patches·m−1 and a density ranging between 28 and 204 colonies·m−2, with a fairly continuous bathymetric distribution. The highest red coral densities in the investigated areas were found on cliffs and boulders mainly exposed to the east, at the greatest depth, and characterized by medium percentage sediment cover. The study populations contained a high percentage (46% on average) of harvestable colonies (>7 mm basal diameter). Moreover, some colonies with fifth-order branches were also observed, highlighting the probable older age of some components of these populations. The Ischia population showed the highest colony occupancy, density and size, suggesting a better conservation status than the populations at the other study locations. These results indicate that deep dwelling red coral populations in non-stressed or less-harvested areas may diverge from the inverse size-density relationship previously observed in red coral populations with increasing depth. [ABSTRACT FROM AUTHOR]

Deep-sea reef-building corals are among the most conspicuous invertebrates inhabiting the hard-bottom habitats worldwide and are particularly susceptible to human threats. The precious red coral ( Corallium rubrum, L. 1758) has a wide bathymetric distribution, from shallow up to 800 m depth, and represents a key species in the Mediterranean mesophotic reefs. Several studies have investigated genetic variability in shallow-water red coral populations, while geographic patterns in mesophotic habitats are largely unknown. This study investigated genetic variability of C. rubrum populations dwelling between 55 and 120 m depth, from the Ligurian to the Ionian Sea along about 1500 km of coastline. A total of 18 deep rocky banks were sampled. Colonies were analyzed by means of a set of microsatellite loci and the putative control region of the mitochondrial DNA. Collected data were compared with previous studies. Both types of molecular markers showed high genetic similarity between populations within the northern (Ligurian Sea and Tuscan Archipelago) and the southern (Tyrrhenian and Ionian seas) study areas. Variability in habitat features between the sampling sites did not affect the genetic variability of the populations. Conversely, the patchy distribution of suitable habitats affected populations' connectivity within and among deep coral banks. Based on these results and due to the emphasis on red coral protection in the Mediterranean Sea by international institutions, red coral could be promoted as a 'focal species' to develop management plans for the conservation of deep coralligenous reefs, a reservoir of marine biodiversity. [ABSTRACT FROM AUTHOR]